United States
Environmental Protection
Agency
Environmental Monitoring and Support— EPA-600/4-78-060
Laboratory         October 1978
Cincinnati'OH 45268
Research and Development
A Manual for the
Identification of the
Larvae of the Caddisf ly
Genera Hydropsyche
Pictet and Symphitopsyche
Ulmer  in Eastern and
Central North America
(Trichoptera:Hydropsychidae)

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r
                                           RESEARCH REPORTING SERIES

                           Research reports of the Office of Research and Development, U.S. Environmental
                           Protection Agency, have been grouped into nine series. These nine broad cate-
                           gories were established to facilitate further development and application of en-
                           vironmental technology. Elimination of traditional grouping was consciously
                           planned to foster technology transfer and a maximum interface in related fields.
                           The nine series are:

                                 1.  Environmental Health  Effects Research
                                 2.  Environmental Protection Technology
                                 3.  Ecological Research
                                 4.  Environmental Monitoring
                                 5.  Socioeconomic Environmental Studies
                                 6.  Scientific and Technical Assessment Reports (STAR)
                                 7.  Interagency Energy-Environment Research and Development
                                 8.  "Special" Reports
                                 9.  Miscellaneous Reports

                           This report has been assigned to the ENVIRONMENTAL MONITORING series.
                           This series describes research conducted to develop new or improved methods
                           and instrumentation for the identification and quantification of environmental
                           pollutants at the lowest conceivably significant concentrations. It also includes
                           studies to determine the ambient concentrations of pollutants in the 'environment
                           and/or the variance of pollutants as a function of time or meteorological factors.
                           This document is available to the public through the NationaJ Technical Informa-
                           tion Service, Springfield, Virginia 22161.

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                                                              EPA-600/4-78-060
                                                              October 1978
    A MANUAL FOR THE IDENTIFICATION OF THE LARVAE OF THE CADDISFLY GENERA
HIDROPSICHE PICTET AND SIMPEITOPSJCEE ULMER IN EASTERN AND CENTRAL NORTH AMERICA
                         (TRICHOPTERA:HYDROPSYCHIDAE)
                                     by
                             Guenter A.  Schuster
                      State Biological Survey of Kansas
                          The University of Kansas
                          Lawrence,  Kansas 66044

                                     and

                               David A-  Etnier
                              Dept.  of Zoology
                         The University  of Tennesee
                         Knoxville,  Tennessee 37916
                              Project Officer

                              Donald J. Klemm
                          Aquatic Biology Section
             Environmental Monitoring and Support Laboratory
                           Cincinnati, Ohio  45268
              ENVIRONMENTAL MONITORING AND SUPPORT LABORATORY
                    OFFICE OF RESEARCH AND DEVELOPMENT
                  U.S. ENVIRONMENTAL PROTECTION AGENCY
                        CINCINNATI,  OHIO  45268

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                                 DISCLAIMER
     This report has been reviewed by the Environmental Monitoring and
Support Laboratory, U.S.; Environmental Protection Agency,  and  approved
for publication.  Approval does not signify that the contents  necessarily
reflect the views and policies of the U.S. Environmental Protection Agency,
    does mention of trade names or commerical products constitute endorse-
nor
ment or recommendation for use.
                                       4

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                                FOREWORD

     Environmental measurements are required to determine the quality of
ambient water, the character of effluents, and the effects of pollutants
on aquatic life.  The Environmental Monitoring and Support Laboratory -
Cincinnati conducts research to develop, evaluate, and promulgate methods
to:

     *  Measure the presence and concentration of physical, chemical,
        and radiological pollutants in water, wastewater, bottom sediments,
        and solid waste.
     *  Concentrate, recover, and identify enteric viruses, bacteria, and
        other microorganisms in water.
     *  Measure the effects of pollution on freshwater, estuarine, and
        marine organisms, including the phytoplankton, zooplankton,
        periphyton, macrophyton, macroinvertebrates, and fish.
     *  Automate the measurement of physical, chemical, and biological
        quality of water.
     *  Conduct an Agency-wide quality assurance program to assure
        standardization and quality control of systems for monitoring
        water and wastewater.

     The effectiveness of measures taken to protect the biological
integrity of the Nation's surface waters is dependent upon our knowledge
of the environmental requirements of aquatic organisms and our understand-
ing of the the complex relationships that prevail in aquatic ecosystems.
Caddisflies are important components of aquatic food webs and are useful
water quality indicator organisms.  This manual contains an illustrated
key and descriptions for 39 species of caddisfly larvae, previously difficult
to identify to the species level.  Also included are notes on the ecology
and distribution of each species, intraspecific variation, synonomies, and
a complete literature survey.  The manual was developed to assist
biologists in evaluating data collected during studies of the effects of
toxic substances and other pollutants on the structure of indigenous
communities of aquatic organisms.
                                           Dwight G. Ballinger
                                           Director
                                           Environmental Monitoring and
                                           Support Laboratory - Cincinnati
                                    iii

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                                  ABSTRACT

     Larvae of the caddisf ly genera Hydrops-yoke and Symphitopsyohe are among
the most encountered and abundant organisms of lotic environments in eastern
North America.  Yet, little is known of the larval stages of these genera.
Previously, the larvae of only 12 species, of which descriptions are presented
here, were known.  Descriptions of larvae of an additional 27 species are here
presented for the first time.

     Larval-adult associations were made by simultaneously collecting metamor-
photypes (pharate adults) and larvae.  Species determination is based on the
cleared male genitalia of the pharate adults.  The abdomens were cleared in
strong KOH so that sclerotized structures lying beneath surrounding tissues
could be examined.  Larvae were studied using both compound and dissecting
microscopes; scale hairs, club hairs, and minute spines of the abdomen are
best studied under high magnification.  The larval abdomens were cleared in
strong KOH and put into glycerin on a microscope slide for examination.

     Presented here are the descriptions of the larvae of 14 of 15 nominal
eastern species of the genus Symphitopsyahej and 25 of 34 of the genus Hydro-
psyche; 18 of 24 of the scalavi,s group, 6 of 9 of the depTavata group, and
the single species of the ouanis group, H. auanis Ross.  A key is provided
for known larvae, incorporating a number of characters previously unused in
the taxonomy of these two genera.  The key is based on ultimate or penultimate
larval instars since color patterns may be more variable in earlier instars.

    Larvae of S. piatrix have not definitely been associated with metamorpho-
types; however, larvae were collected near the type locality, and the presumed
larvae of this species is described, illustrated, and keyed.  A listing of all
unassociated species, with known distribution and literature citations, is
given.

     Characters utilized in the key  to facilitate separation of species
include:  color pattern of head; morphology of anterior margin of frontocly-
peus; presence of large tubercle on  center of anterior ventral apotome; pre-
sence or absence of club hairs, scale hairs and/or minute spines on dorsum of
abdomen; and presence of large, heavily sclerotized spine-like setae on venter
of anal legs.  Minute abdominal spines are described and utilized as a taxo-
nomic character for the first time.

     In addition to the description  of the larvae,  the following are presented
for  each of the associated species of Hy dropsyahe and Symphitopsyohe:  known
range, notes  on  the biology, diagnosis, intraspecific variation, material
examined,  complete  literature survey and  synonomies, and illustration  of  the
head capsule  and pronotum.
                                       IV

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                                  CONTENTS

Foreword	    ill
Abstract	     iv
Figures	    vli
Acknowledgment	     xi

Section                                                                  Page

  1. Introduction  	      1

  2. The Geographic Scope   	      4

  3. Collecting Techniques	      5

  4. Methods	      7

  5. Material Examined and  Borrowed	     10

  6. Generalized Description of Hydvopsyohe
          and Symphitopsyohe Larvae	     11

  7. Key to the Known Larvae of the Genera Hydropsy che
          and Symphitopsyohe in Eastern North America	     13

  8. Discussion of Species
          The Genus Symphitopsyohe
               Symphitopsyohe b-Lfida  (Banks)	     30
               Symphitopsyohe dheilon-is (Ross)	     33
               Symphitopsyohe Teouwata (Banks)	     34
               Symphitopsyohe walkevi (Betten and Mosely)	     35
               Symphitopsyohe bronta  (Ross)	     37
               Symph-Ltopsyehe movosa  (Hagen)	     41
               Symph-Ltopsyehe riola  (Denning)	     44
               Symphitopsyche alhedra (Ross)  	     45
               Symphitopsyohe slossonae (Banks)   ...  	     47
               Symphitopsyohe maoleod-i  (Flint)	     50
               Symphitopsyohe spccma  (Ross)	     52
               Symphitopsyohe ventuTa (Ross)  	     55
               Symph-i-topsyohe piatrix (Ross)	     57
               Symphitopsyohe etnievi (Schuster and  Talak)  	     58
          The Genus Hydropsyohe
          Depx>avata Species Group
               Hydropsyohe  betteni Ross	     61
               Eydropsyohe  depravata  Hagen  	     63
                                      v

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Section                                                                  Page
               Eydropsyohe potomacens'i.s Flint	     64
               Eydropsyohe el-lssoma Ross	     65
               Eydropsyohe deodlda Ross	     66
               Eydropsyohe Carolina Banks	     68
          Cuccnis Species Group
               Eydropsyohe cuanis Ross	     70
          Saalaris Species Group
               Eydropsyohe orris Ross  	 .....     71
               Eydropsyohe Widens Ross	     75
               Eydropsyohe aerata Ross 	     77
               Eydropsyohe phalerata Hagen  	     78
               Eydropsyohe dicantha Ross	     80
               Eydropsyohe demora Ross 	     83
               Eydropsyohe valanis Ross	     85
               Eydropsyohe arinale Ross	     86
               Eydropsyohe soalari-s Hagen	     87
               Eydropsyohe s-imulans Ross	     90
               Eydropsyche inoonmoda Hagen  	     92
               Eydropsyohe frison-i Ross	     93
               Eydropsyohe miss-iss-ippiensi-s Flint	     95
               Eydropsyohe venular-is Banks	     96
               Eydropsyohe hoffmani, Ross	     98
               Eydropsyohe leonardi Ross	    100
               Eydropsyohe hagen-i Banks	    102
               Eydropsyohe patera Schuster and Etnier   	    104

Literature Cited 	    107

Glossary	    116

Appendixes
     Appendix A.  Species Associated From Each
          Genus and Species Group With a Listing
          of States From Which the Metamorphotypes
          and/or Larvae Were Examined  	    119
     Appendix B.  Unassociated Eydropsyohe and
          Symphitopsyche Species of Eastern North
          America With Listing of All Literature
          and Known Distribution	    124
                                     vi

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                                   FIGURES
Figure
Page
  1.  Hydropsyohe k>etteni Larva,  Lateral Aspect
           (Adapted and Modified from Ross, 1944)  	     19

  2.  Hydropsyche simulans Larva, Head, Ventral
           Aspect	     20

  3.  Hydropsyche simulans Larva, Head, Dorsal
           Aspect	     20

  4.  Hydropsyche simulans Larva, Labrum, Dorsal
           Aspect	     20

  5.  Hydropsyche simulans Larva, Mandibles,
           Dorsal Aspect 	     20

  6.  Hydropsyshe simulans Larva, Prosternum and
           Poststernal Plates  	     20

  7.  Hydropsyohe sp. Pupa, Case, Dorsal Aspect	     21

  8.  Hydropsyche sp. Pupa, Case, Ventral Aspect 	     21

  9.  Hydropsyche simulans Larva, Apex of Abdomen,
           Dorsal Aspect (Modified from Ross, 1944)  	     23

 10.  Hydropsyche simulans Larva, Abdominal
           Segment VII, Dorsal Aspect, Enlarged
           Portion of Epidermis (After Ross, 1944) 	     23

 11.  Hydropsyche arinale Larva, Abdominal Segment
           VII, Dorsal Aspect, Enlarged Portion of
           Epidermis (After Ross, 1944)  	     23

 12.  Symphitopsyohe recurvata Larva, Abdominal
           Segment VII, Dorsal Aspect, Enlarged
           Portion of Epidermis (After Ross, 1944) 	     23

 13.  Hydropsyche dicantha Larva, Head, Lateral
           Aspect; Arrow Shows Bristle-Like Setae
           on Genae and Frontoclypeus	     23
                                     vii

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14.  Hydeopsyche aevata Larva, Apex of Abdomen,
          Ventral Aspect (Inset showing minute
          spines at base of stout setae)  (After
          Ross, 1944)	     23

15.  Hydvopsyche sp. Larva, Frontoclypeus; A.
          Dorsal Aspect Showing Small Spine-like
          Setae on Posterior Portion of Sclerite;
          B.  Lateral Aspect	     23

16.  Hydropsyche hoffmani Larva, Head, Lateral •
          Aspect; Arrow Shows Tubercle on Anterior
          Ventral Apotome .	     23

17.  Symphitopsyche biftda Larva, Head, Dorsal
          Aspect	     25

18.  Symphitopsyche walkeri Variant Larva, Head,
          Dorsal Aspect 	     25

19.  Symphitopsyohe bvonta (Appalachian Form)
          Larva, Head, Dorsal Aspect  	 	     25

20.  SympTritopsyche movosa Larva, Head, Dorsal
          Aspect	     25

21.  Symphi-topsyche riola Larva, Head, Dorsal
          Aspect	     25

22.  Symphitopsyohe alhedra Larva, Head, Dorsal
          Aspect	     25

23.  Symphi-topsyche slossonae Larva, Head, Dorsal
          Aspect	     25

24.  Symphitopsyohe slossonae Variant Larva, Head,
          Dorsal Aspect 	     25

25.  Symph-i-topsyohe maoteodi Larva, Head, Dorsal
          Aspect	     25

26.  Symphitopsyche sparna Larva, Head, Dorsal
          Aspect	     25

27.  Symph'i-bopsyche ventwva Larva, Head, Dorsal,
          Aspect	     25

28.  SymphLtopsyehe ptatr-ix Presumed Larva, Head,
          Dorsal Aspect		     25

29.  Sympkltopsyche etnlevi. Larva, Head, Dorsal
          Aspect	••     25
                                   viii

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30.  Hydropsyohe betteni Larva, Head, Dorsal
          Aspect  	
                                                                         25
31.
32.


33.


34.


35.


36.


37.


38.


39.


40.


41.


42.


43.


44.


45.


46.
     Hydropsyohe el-Lssoma Larva, Head; A.
          Dorsal Aspect; B.  Lateral Aspect,
          Arrow Showing Rounded Tubercle at
          Posterior Angle of Frontoclypeus  .

     Hydropsyohe deoalda Larva, Head, Dorsal
          Aspect  	
     Hydropsyohe carol-ina Larva, Head; A.
          Dorsal Aspect;  B.  Dorsolateral Aspect

     Hydropsyohe ouanis Larva,  Head, Dorsal
          Aspect (After Ross, 1944)  	
     Hydropsyohe orris Larva, Head; A.   Dorsal
          Aspect; B.   Lateral Aspect  .  .  .  .  .
     Hydropsoyhe bidens Larva,  Head;  A.   Dorsal
          Aspect; B.   Lateral Aspect   	
     Hydropsyohe aerata Larva,  Head,  Dorsal
          Aspect  	
     Hydropsyohe phalerata Larva,  Head,  Dorsal
          Aspect  	
     Hydropsoyhe dioantha Larva,  Head,  Dorsal
          Aspect  	
     Hydropsoyhe demora Larva,  Head,  Dorsal
          Aspect	f	
     Hydropsyohe valon-Ls Larva,  Head,  Dorsal
          Aspect  	
     Hydropsyohe arinale Larva,  Head,  Dorsal
          Aspect (After Ross,  1944)  .  .  .  .  .
     Hydropsyehe  soalaris Larva,  Head,  Dorsal
          Aspect   	  	
     Hydropsyohe  simulans Larva,  Head;  A.
          Dorsal  Aspect;  B.   Lateral Aspect
     Eydropsyche mississ-ippiensis Larva,  Head,
          Dorsal Aspect •  •  •  •	
     Hydropsyahe venularis  Larva,  Head;  A.
          Dorsal Aspect;  B.   Lateral Aspect
25


27


27


27


27


27


27


27


27


27


27


27


27


29


29


29

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47.  Hydropsyche hoffmani Larva, Head, Dorsal
          Aspect  	
48.  Hydropsydhe leoncccdi. Larva, Head; A.
          Dorsal Aspect; B.  Lateral Aspect

49.  Hydropsyche hageni Larva, Head; A.
          Dorsal Aspect; B.  Lateral Aspect

50.  Hydpopsyehe patera Larva, Head,
          Dorsal Aspect •  •  •  •  •  v •  •  •  *
51.  Hydeopsyehe fr-ison-C Larva, Head; A.
          Dorsal Aspect .	
29
29
29
29
29

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                               ACKNOWLEDGMENTS

     This publication  is adapted,  in part,  from a dissertation presented to
 the  faculty of  the Graduate  School of the University of Tennessee by the
 senior author,  in partial  fulfillment of the requirements for the degree of
 Doctor of Philosophy.

     We are very appreciative of the role the State Biological Survey of
 Kansas has played in the publication of this manuscript.  We are grateful to
 its  director, Dr. Ronald L.  McGregor, and to Donald G. Huggins, in charge of
 the  Aquatic Invertebrate Survey Section, for their encouragement and support,
 and  especially  for providing the clerical assistance necessary to prepare the
 photo-ready copy of this manuscript.

     The senior author would like  to express his appreciation to Drs. Dewey
 Bunting, Melbourne Whiteside, and  Susan Riechert, Department of Zoology, and
 Dr.  Paul Parmalee, Department of Anthropology, for their criticisms of the
 manuscript and  for serving on his  doctoral  committee.

     A special  expression of gratitude is extended to Drs. Oliver S. Flint,
 Jr., Department of Entomology, Smithsonian  Institute, and John D. Unzicker,
 Faunistics Survey Section, Illinois Natural History Survey, for freely giving
 much valuable time and advice during the course of this study.  We are also
 grateful to them for allowing the  senior author to visit their respective
 institutions, study specimens in their holdings, and borrow extensive amounts
 of Hydropsyche and Symphi-topsyche material.

     We wish to express gratitude to the many people who contributed  speci-
 mens or assisted in collections for this study, especially the following:
 Fae Andrews, Noel Burkhead, William Dickinson, Mark Hughes, Lynn and Wayne
 Starnes, Anthony Talak, Virginia Tolbert, G. William Wolfe.

     Special consideration is given to the following who gratiously supplied
 larvae and/or metamorphotypes of Hydropsyohe and Symphitopsyche species for
 study:  Dr. Bruce Wallace, University of Georgia (5.  maoleodV);  Dr.  Kenneth
 Stewart, North Texas State University (E.  simulans);  and Dr. Todd Harris,
 Purdue University (5.  spccma, H.  betteni,  and S.  morosa);  Drs. G. L. Harp,
Arkansas State University and H.  W. Robinson,  Southern Arkansas University
 (presumed larva of 5.  p-Laiwix) .

     Gratitude is also expressed to Sigma Xi for awarding a grant which made
 possible several fruitful collecting trips and to Highlands Biological Sta-
 tion, Highlands, North Carolina,  for supporting two  weeks  of study at the
 Biological Station;  and to the Southern Region Educational Board for the
 funding which made the visits to the Smithsonian Institute and Illinois
Natural History Survey possible.

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     Special gratitude is given to Dr. H. H. Ross for allowing us to redraw
illustrations which appeared in his 1944 work, "The Caddis Flies, or
Trichoptera, of Illinois."  We would also like to express our appreciation
to Drs. Oliver S. Flint* Jr., Andrew P. Nimmo and Glenn B. Wiggins for re-
viewing this manuscript and offering valuable suggestions which considerably
improved the final product.

     We are grateful to Ms. Claire Schuster for typing the first draft of the
manuscript, and to Ms* Jan Powers for typing the final camera-ready copy.
Both attacked this chore with great enthusiasm even after numerous retyping
of pages.

     Last, and most of all, we wish to thank our wives and families for their
assistance in the field, and their support throughout this study.
                                  - xii

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                                   SECTION 1

                                 INTRODUCTION

      Studies of North American Trichoptera have been most concerned with adult
 systematics (Betten,  1934;  Ross, 1944).   In general, it is true that the
 systematics of adult  caddisflies is much better understood than that of their
 corresponding larval  forms.  Of the nearly 1,000 described Trichoptera in
 North America, the immature stages of only 20 percent of these species are
 known (Wiggins,  1964).

      Resh (1973:1)  points out that "the  need for keys to identify the immature
 stages of caddisflies has become increasingly important with regard to under-
 standing  their potential as water quality indicators.   Very common genera,
 such  as Cheumatopsyche  and  Hydropsyche (sensu Ross,  1944)  of the family
 Hydropsychidae are almost totally unknown in the aquatic stages."  Studies  on
 caddisfly larval taxonomy are not only important for the realization of their
 potential as water quality  indicator organisms but also, are vital for a number
 of other  reasons.

      First,  such studies may be used as  a starting point from which natural
 history and a wide spectrum of ecological researches may be based.   Any such
 studies hinge on correct species determination.   Second,  and most important
 from  the  "trichopterologist's" point of  view,  such studies  give  a new dimen-
 sion,  the larva,  from which data may be  gleaned.  A  knowledge of the immature
 stages  may develop  a  greater understanding of  the systematics and phylogenetic
 relationships  within  Trichoptera.   Ultimately,  these may give rise to  a better
 understanding  of  the  entire order Trichoptera.

      Early works  on North American Trichoptera  (Say, 1823,  1824,  1828; Walker,
 1852; Hagen,  1860,  1861, 1864,  1866,  1868, 1873,  1875; McLachlan,  1863;  Banks,
 1894, 1895,  1897, 1899, 1900a,  1900b, 1900c, 1901a,  1901b,  1903a,  1903b,
 1904a,  1904b,  1904c,  1904d,  1905a,  1905b,  1907a,  1907b,  1908a, 1908b,  1909,
 1911, 1914,  1916, 1918, 1920,  1924,  1930a, 1930b, 1936a, 1936b,  1938,  1943,
 1944; Betten,  1950; L. Milne,  1934-36) were almost completely restricted to
 adult systematics.  It was not until  the  turn of  the century  that workers
 began to  study the  larvae in  greater  detail.  Workers such  as  Betten  (1902,
 1934), Lloyd  (1915, 1921), M. Milne  (1938, 1939)  laid the foundations  for
 early larval work.  However,  it was not until the classic work by Ross  (1944),
 "The Caddis Flies, or Trichoptera of  Illinois", was published  that taxonomic
keys were available for the greater majority of Trichoptera larvae.  This
work has been the single most authoritative study on North American caddis-
flies.  It not only dealt with the species of Illinois, but also most amply
illustrates all known widespread species.  In it Ross compiled keys to the
larvae of families and genera and, whenever known, to the species level.  The
importance of this work cannot be overstressed.

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     Wiggins (1977) is the major contribution dealing with larval systematic^,
on the generic level, for the entire order since Ross, 1944.  This work is an
indicator that in recent years researchers have increasingly directed their
attention to larval forms.  The systematics of immature forms has been worked
out for a number of genera and families.  Wiggins (1960) discussed the larvae
of the caddisfly family Phryganeidae; Wiggins and Anderson (1968) made special
reference to the immature stages of Pseudostenophylax and Ph-ilocasca.  Wiggins
(1973a, 1973b) discussed the larval systematics of several genera of the
family Limnephilidae.  Merrill and Wiggins (1971) and Yamomoto ^and Wiggins
(1964) respectively described the larvae of Setodes and Mystacides of the
Leptoceridae.  Other authors have worked extensively with larval taxonomy;
Flint (1960), the taxonomy and biology of limnephilid larvae; Flint  (1961),
the Immature stages of Arctopsychinae; Flint (1962), Rhyacophila; Flint (1964),
Psychomyiidae; Wallace and Sherberger  (1970) , Calamoceratidae; Sherberger and
Wallace (1971), Molanna.', Wallace (1971), Brachycentrus', Ross and Wallace
(1974), Sericostomatidae; Resh  (1973, 1976), Ceraclea.

     Recent European authors have also done a great deal of larval work;
Lepneva (1964, 1966), the larvae and  pupae of Annulipalpia and Integripalpia,
respectively; Hicken (1967), the caddis larvae of Great Britain; Hildrew and
Morgan (1974), the taxonomy of British Hydropsychidae; Sed Lak (1971),
Hydropsyche of CheckoSlovakia.

     The genera Hydropsyche and Symphitopsydhe consist of approximately 70
species in North America of which about 50 are restricted to eastern and cen-
tral North America.   The North American species of Hydropsyche  are  divided
into three species groups  (Ross, 1944);  the cuanis group,  the depravata
group, and the scalaris group.  The  cuanis group consists of a single species,
ff. cuanis Ross.  The depravcrta  group  consists of nine nominal species,^all of
which are found in eastern and  central North America  with H. guttata Pictet
being holarctic in distribution.  The remaining  32  species  belong  to the
scalaris group of  which 24 are more  or  less restricted  to eastern  and central
North America.

     The genus Symphitopsyche was until recently (Ross  and  Unzicker, 1977)
considered part of Hydropsyche  (the  bifida species  group  in Ross,  1944).  The
genera as adults are separable  primarily  on the  basis of  the morphology of
the  aedeagus and the 9thiand  10th abdominal segments.  As larvae they are
separable on the basis  of club  and  scale  hairs and  absence  or presence of
minute spines on the dorsum of  the  abdomen.  Symphitopsyche in North America
is composed of approximately  30 species of which about half are  found  in
eastern  and central  North America.

      The only key  to the larvae of these  two  genera is that of Ross (1944)  in
which both genera  were treated  under Hydropsyche.   Twelve species are keyed
and 2 additional species, not keyed, are  briefly described.  The study of the
 larval taxonomy of the g'enera Hydropsyche and Symphitopsyche has been in
 limbo since that publication,  even though the larvae are commonly encountered,
 often in very large numbers in almost all types  of lotic environments in
 eastern North America.                                i     '.  '

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     The primary purpose of this study is to associate the larvae of the
species of HydropsyaTie and Symphitospydhe distributed in eastern and central
North America and to construct a key for the identification of these larvae.
Secondary reasons for this study are:  (1) to present distributional data on
many of the little known species in these genera, (2) to give emergence dates
to assist future workers of Hydropsyche and Symphitopsyche,. and (3) to give
notes on the biology of all species that have been associated.

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                                  SECTION 2

                            THE GEOGRAPHIC LIMITS

     At the onset of this study, it was determined that the southeastern
United States be used as the study area.  However, as the study progressed,
an increasing number of species were associated so that it became feasible to
encompass the entire eastern and central North American region.

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                                   SECTION 3

                             COLLECTING TECHNIQUES

      Collections were made in springs, streams,  and rivers;  all types of lotic
 environments were sampled.  A collecting site was selected on the basis of the
 amount of riffle habitat in the area.   Hydropsyehe and Symphitopsyohe larvae
 prefer these types of habitats over areas consisting of a long, smooth run.
 Each likely site was systematically examined for larvae and metamorphotypes
 (M.  Milne*  1934; pharate adults,  sensu Hinton, 1971 and Wiggins,  1977).
 Initially,  at each locality,  the fastest water was selected to be sampled
 first.   In each riffle,  rocks were picked up and examined for larvae and
 pupae.   Since some Hydropsyehe and SymphLtopsyche species seem to have a
 definite preference for  rock size and  shape,  many types and  shapes of rocks
 were examined.   Often larvae and  pupae may congregate in spaces between
 rubble on the bottom.  These areas were also examined.   When the  boulders
 were too large to move,  they were examined underwater.   While steadily
 anchoring oneself in the riffle or rapid,  a hand may be taken and gently run
 across  the  surfaces and  crevices  of the rocks.   With some experience one
 obtains a feeling for larval  nets and  pupal cases,  and  these may  be grasped
 underwater.   Although this is a tedious task^ it is a very fruitful method
 with strata which cannot otherwise be  sampled.

      Once the riffle habitats were thoroughly examined,  other areas of  the
 stream  or river were then collected.   Runs  may often be more difficult  to
 collect than  riffle areas  because they are  usually much deeper, and submerged
 rocks,  logs,  or branches are  not  as  accessible.   Long-handled aquatic nets
 and  dredges are of  use in  such areas.

      Sand streams and streams  below  the fall  line also  presented  collecting
 problems.  In such  streams, which may  be sluggish,  it was  often difficult to
 determine where the Hydropsyche habitat existed.   It was often exasperating
 to collect Hydropsyche adults  in  such  areas and not  find any  larvae or pupae.
 It was  found  profitable, while  collecting such streams,  to put submerged
 materials such as debris, Iqgs, and branches  in the  sun  to dry.  Larvae which
 had burrowed  into these ^o.u^d  invariably crawl out  in a  few minutes and could
 then  easily be collected.,.'-Collecting metamorphotypes here, however, was not
 so easy.  To  obtain  these, jthe  submerged materials were picked or torn apart
 piece by  piece after the ^aryae were,collected.  This was very time consuming.
 If time was of the  essence, ''these materials were put into the collecting
vehicle and picked enroute to the next site or were examined later that
 evening.

     At each locality an effort was made to obtain adults by searching emer-
gent and marginal vegetation.  The undersides of bridges were also examined.
During the day, adults were captured with the use of an aspirator.  In the

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evening collections were made in a number of ways.  Firstly by examining neon
signs and other lights near the stream; secondly, portable light traps were
set up along the banks of the stream.  These lights were either flourescent
or blacklights powered by 6 volt batteries.  Placed near each light was_a pan
half filled with 70% isopropyl alcohol, and adults would drop directly into
these pans.  Third, when flourescent or blacklights were unavailable, a   _
Coleman lantern was used in a similar manner as described above.  In addition
to collecting adults directly into alcohol pans, it was convenient to place
the lights near a draped white sheet  or  the hood of a car; adults were
aspirated as they alighted on these.  This method allowed more selectivity in
the forms collected.

     All adults collected:were preserved in 70% to 75% isopropyl alcohol.
The larvae and metamorphotypes were  initially collected in vials containing
95% to 100% isopropyl alcohol because of the large amounts of water introduced
into the vials with larvae and pupal cases.  If large quantities of larvae and
pupal cases were collected, alcohol  was changed immediately after the collec-
tion was made and again within 2  or  3 days with 70% isopropyl alcohol after
the specimens had sufficiently hardened.

     Previous published and unpublished Hydropsydhe and Symphitopsyche
adult collection records  contributed greatly to  the success of  this  study.
These records gave  important data with regard to  locality and dates  or
probable emergence  periods.  These provided an  excellent  starting point  for
collecting larvae and metamorphotypes.

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                                  SECTION 4
                                   METHODS

     Initially, it was intended that many species of Hydropsyche and Symphi--
topsyo'he larvae and adults would be associated by larval rearing.  However,
because of the need for constant running water at cold temperatures and the
existence of probably a number of limiting factors such as dissolved oxygen,
photo-period, suspended nutrients, and various chemical requirements not yet
defined, our limited attempts at rearing met with failure.  Therefore, this
method was abandoned so that all the investigative efforts and time could be
employed collecting metamorphotypes (mmts.).  The "metamorphotype method" was
first described by M. Milne (1934), and proved invaluable in this study.

     The metamorphotype or pharate adult is a feature of great importance to
the study of Trichbptera.  It represents a period in the life of the indivi-
dual caddisfly in which all three stages of the holometabolous life cycle
are available for study — larva, pupa, and developing adult.

     The larva encapsulates itself into a silken case (Fig. 7-8) to the out-
side of which rocks, sand, vegetation, or other debris is glued.  The under-
side is shaped by the substrate.  As the larva undergoes metamorphosis and
changes into a pupa, the larval sclerites are sloughed and packed into the
back of the case where they remain during subsequent development.  The pupa
then develops into the adult with the formation of wings, partial atrophy of
mouth parts, and sclerotization of genitalia.  If these are collected after
the sclerotization of genitalia and before emergence of the adult, the inves-
tigator has available the larval sclerites, pupal skin, and adult genitalia
by which species identification is made.  Therefore, an absolute larval-adult
association has been accomplished.

     During this study, it was found that one may reliably tell if the adult
genitalia were sufficiently sclerotized by the color of the eyes of the pupa.
If the eyes were dark rather than white, one could be assured of sufficient
sclerotization.  Dark wing coloration may also be a good indicator of this.
If only immature pupae are collected, they may be placed into small jars, on
moist sphagnum moss, and stored in a refrigerator until further development
has taken place (Wallace, personal communication).

     Sclerotization of genitalia is important for several reasons.  First,
soft genitalia may adhere to the pupal skin and make adequate examination
impossible.  Second, the true shape of the aedeagus and claspers is not
evident until sclerotization has taken place.  For this reason examination of
an immature specimen may lead to faulty identification if extreme care is not
taken.  Last, it is often necessary to examine morphological features of the

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genitalia which are imbedded in. surrounding tissue.   To do this,  the abdomen
is cleared in solution of KOH.  If the genitalia are,not adequately sclero-
tized, some of the features may be lost by the action of the KOH.

     Once mature pupae were collected, preserved, and hardened,  they were
examined and the larval sclerites retrieved from the pupal case.   Hydropsyche
and Symphitopsyehe pupal cases usually were attached to rocks or  other sub-
strates.  Because of this, one side of the case is flat and without an outer
covering of granules in order to attach snuggly to the substratum.  This pro-
vides an excellent area to open the case and assure minimal damage to the
pupa and little loss of the larval sclerites.  Often,  however, Hydropsyahe
and Symphitopsyehe pupal cases may be attached to strands of Potamogeton or
Podostemm.  If so, the pupal case was completely cylindrical with granules
(Wentworth Scale, after Wentworth, 1922) attached on all sides.   It was
extremely hard to tell in such situations which was anterior or  posterior.
One must take great care in opening these cases since the granules were
usually rigidly attached ;to one another and difficult to pry apart.  It was
best to begin prying near either end of the case.  With the removal of a few
granules, one can quickly observe either the anterior or posterior end of the
pupa.  If the posterior end was opened, it was found best to stop and open the
other end.  This prevented scattering of larval sclerites.  Once the anterior
end has been opened, very fine forceps may be used to extract the pupa from
the case.  After the pupa has been removed undamaged,  the rest of the case may
be opened to retrieve the larval sclerites.

     The larval sclerites were picked up individually and placed into a
microvial.  After all the larval sclerites were placed in the microvial, the
pupa was also put into it, thereby isolating each pupa and its sclerites from
those of other specimens.

     The microvials are of clear plastic tubing with outer diameter of 4.0 mm
and inner diameter of 3.5 mm.  The tubing was cut into 15 mm sections with one
end being heated and pinched off.  These made excellent microvials as pupae of
most Eydropsyche and Symphitopsyehe species fit snuggly into them, and cotton
stoppers were not required.  This allowed for storage of 30 to 40 pupae and
sclerites in one 3-dram vial.

     Since not all individuals in a Hydropsyche or Symphi-topsyohe population
pupate simultaneously, larvae of the same species were usually collected with
metamorphotypes.  Larval sclerites from the metamorphotypes collected were
matched with larvae collected at the same locality.  This resulted in defi-
nite larval-adult associations.  If no larvae were collected with the meta-
morphotypes, attempts were made to go back at a later time specifically to
collect larvae.

     The larva of S. piatrix  (Ross) has not definitely been associated using
metamorphotypes.  However, larvae were collected near the type locality, and
the lack of other nonrecognizable larvae presents strong evidence that these
larvae are in actuality those of the above species.  The presumed larva of
this species is described, illustrated, and keyed.

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     Species identification of metamorphotypes was based on genitalia.  Present
adult Eydropsyohe and Symph-i.topsyahe taxonomy is based solely on the genitalia.
The pupal skin was carefully removed from the abdomen to minimize the damage.
This exposed the genitalia for close observations.  Often the abdomen had to
be cleared for examination of internal structures embedded in surrounding
tissues.

     The clearing process was done as described by Nimmo (1971).   A strong
solution of potassium hydroxide was made, into which the detached pupal abdo-
men was submerged.  This solution was brought to a gentle boil for a few
minutes.  The abdomen was then retrieved and placed in a weak solution of
acetic acid (10%) and returned to the hot KOH.  This produced a massive re-
lease of bubbles which dislodged tissue surrounding the genitalia.  This pro-
cess was repeated until all surrounding tissue was cleared.  When this was
accomplished, the abdomen was washed with the acetic acid one last time to
stop the clearing process.  The abdomen may then be placed in glycerin or
alcohol for further examination.  The amount of time required for clearing
could not be standardized.  The size of the specimen and its state of preser-
vation determined the amount of time needed for the process.

     In order that the scale hairs (Fig. 10), club hairs (Fig. 12), and minute
spines (Fig. 10) on the abdomen could be more easily studied, the abdomen of
the larva was removed and treated in a similar manner.  Once the abdomen has
been cleared, it may be put into glycerin on a microscope slide and observed
under a compound microscope.  If the larva was not well preserved, the inter-
nal tissues may be gently teased from the abdomen with very fine forceps; the
end result is the same as that of clearing in KOH.

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                                  SECTION 5

                       MATERIAL EXAMINED AND BORROWED

     Larvae and pupae of the following species were borrowed from the Illinois
Natural History Survey (INHS):  H. arinale, S. bronta, and E. cuan-ls.  Adult
material of H. catawba, H. Iwigen-i., and E. ieonardi were also borrowed from the
INKS.  All other material used to construct keys, and make descriptions, and
illustrations for this study were collected by the authors, or fellow graduate
students at the University of Tennessee, Knoxville.

     All available type material of Eydropsyche and Symphitopsyehe species in
the collections of the INHS and the U. S. National Museum  (USNM) was examined.
Identified material for species, whose types were deposited elsewhere, was
also examined at the above institutions.  A period of 1 week was spent at
each museum by the senior author at which time all species identifications
made during this study x\rere verified.
                                      10

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                                  SECTION 6

                         GENERALIZED DESCRIPTION OF
                    HIDROPSICHE AND SYMPHITOPSICHE LARVAE

     Larva campodieform (Fig. 1).  Head in dorsal view (Fig. 3) subsquadrate
to subrectangular.  Frontoclypeus subtriangular; approximately three—fourths
total length of head; posterolateral margins meet to form an acute
angle; anterior margin straight or convex and toothed.  Labrum (Fig. 4) con-
vex dorsally; anterior margin rounded; outline subtrapazoidal.  Surface of
labrum covered with numerous long setae; lateral margins fringed with long
setal brush.  Mandibles (Fig. 5) triangular with five to seven teeth on apical
half of inner margin; left mandible with tuft of yellow setae above most
proximal tooth; right mandible without such a brush.  Left mandible with large
cavity below dorsal ridge; ridge mandible without such cavity.  Both mandibles
with long, thin setae on lateral margins.  Maxillary palp five segmented;
lacinia conical; hypopharynx mesad of maxillae.  Submentum (Wiggins, 1977)
(Fig. 2) with pair of anterior truncate lobes; anterior ventral apotome
(Wiggins, 1977) (Fig. 2) subtriangular.  Genae fused ventrally with small
triangular posterior ventral apotome or protogula (Fig. 2) (Badcock, 1961) at
posterior end of gular suture.  Ventral surface of genae behind mandible with
band of numerous ridges (stridulatory surface) (Fig. 2).

     All three thoracic segments sclerotized dorsally.  Nota subquadrate;
pronotum with median suture line; mesonotum with large black U-shaped mark
posteriorly; metanotum with posterior short black slash mark; each notum with
black lateral margins.  Strap-like prosternum (Fig.  6) behind first pair of
legs; additional pair of less sclerotized prosternal plates just posterior to
prosternum.  Mesosternum with pair of weakly sclerotized triangular sclerites
behind mesothoracic legs; metasternum lacking sclerites.  Propleuron rectan-
gular with diagonal black line; foretrochantin (Fig. 1) forked with thin,
short, black hairs.  Prothoracic legs more robust, shorter than posterior two;
all segments of forelegs stouter than those of other legs.  Femur of front
legs flattened; posteromesal margin with group of long, heavy, black setae.
Segments of all legs abundantly adorned with long, black setae and short,
golden brown spine-like setae.

     Abdomen covered with many thin, short, black setae.  Scale-like hairs
and minute spines (Fig. 9-10) typically located on dorsum of abdominal seg-
ments of Hydpopsyohe species.  Club-like hairs (Fig. 12) present on dorsum of
abdominal segments of Symphitopsyche species; club hairs as long as other
hairs only slightly thicker; Symphitopsyohe species lack minute spines on
dorsum of abdomen.  Abdominal segments III - VII, respectively, possessing
1, 3, 3, 3, and 2 pleural gills (Fig. 1); these gills are conical, often
covered with short, black setae.  Ventral gills on thoracic segments II - III
                                     11

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and on abdominal segments: I - VII (Fig. 1).  One pair of gills on second
thoracic segment; each of a single stalk with numerous single filaments
attached to it.  Third thoracic segment with four gills; two laterally, two
mesally; filaments similar to these of gills on second segment;  mesal gills
near center of segment.  Gills of abdominal segment I similar to those on
third thoracic segment; medial gills widely separated and close to lateral
gills.  Gills of abdominal segments II - VI similarly arranged;  each segment
with four gills, lateral gills with stalk divided near base; mesal gills con-
sisting of single stalk, not divided.  Each gill with numerous filaments
attached.  Segment VIII of abdomen with two gills; stalk divided at base
with numerous filaments attached.

     Venter of abdominal segments VIII - IX with triangular sclerotized
areas; each sclerite with numerous golden brown stout setae; posterior mar-
gins of sclerites fringed with long, black setae; posteromesal section of
sclerite on segment IX with small triangular area without stout setae.  Sides
of segment IX with pair of smaller, less sclerotized areas with black setae.
Anal legs (Fig. 1) covered with long sclerite dorsally; apical end with large
heavily sclerotized hook; angled ventrally 90 degress; base of hook with
single heavy, long, black seta.  Distal end of dorsal sclerite of anal legs
with fan-like brush of extremely long, black setae; ventral surface of anal
legs membranous and covered with numerous short, black hairs; some species
with heavily sclerotized spine-like setae.  Four anal papillae present
(Fig. 1); may or may not be extruded.
                                      12

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                                 SECTION 7

                     KEY TO KNOWN LARVAE OF THE GENERA
                      HYDROPSYCHE AND SYMPEITOPSYCHE
                         IN EASTERN NORTH AMERICA1

1          Dorsum of abdomen with minute spines on at least
                segments I - III (Fig. 10); scale hairs  (Fig.
                9-11) present on at least the last three
                abdominal segments	Eydvopsyohe .  .	„ 12
1"         Dorsum of abdomen lacking minute spines; club hairs
                (Fig. 12) present on dorsum of abdomen; scale
                hairs lacking	 Symghttopsyohe	2

2,  (I1)    Checkerboard pattern on frontoclypeus (Fig. 17)	f  3
2*         Frontoclypeus with other patterns or unicolored	  4

3  (2)     Posterior angle of frontoclypeus with three distinct
                spots (Fig. 20)	S. movosa (p. 41)
3'         Posterior angle of frontoclypeus with only a single
                large spot* ..... 5. bifida (p. 30); S-. oheilon-iB (p. 33)
                                     S. watkez*i (p. 35); S. reourvata (p. 34)
                                               Central Form S. bronta (p. 37)

4  (21)    Head pattern and thoracic nota black with no distinct
                spots or pattern on frontoclypeus	5
4'         Head and thoracic sclerites yellow to bronze usually
                with some color pattern; if head is black, there
                is at least one central pale spot on the
                frontoclypeus 	  6

5  (4)     Posterior portion of head brown to black with no
                conspicuous circular muscle scars (Fig. 22);
                black hairs and club hairs densely covering
                abdomen; club hairs distinctly thicker than
                regular hairs.  Larva large, robust
S. alhedva (p. 45)
    Based on last instar larvae; color patterns of earlier instars may be
more variable.
   ^
    Scale hairs in H. dioantha may approach the appearance of club hairs,
however this species like all Hydropsyohe, possesses the minute spines on
the dorsum of the abdomen.  The spines under lower magnifications produce
a peppered appearance, and can best be seen on uncleared specimens on the
anterior and posterior margins of the segments.

                                     13

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 6  C4')


 6'


 7  C6')



 7'

 8  (7')



 8f


 9  C8)
10  (8')

10'


11  (10')
11'
Posterior portion of head lighter brown with
     numerous conspicuous circular dark brown
     muscle scars (Fig. 29).  Black hairs and
     club hairs moderately dense on abdomen;
     club hairs only slightly thicker than regular
     hairs. . Larvae not.as large or robust . .  S. etnieri ;(p,. 58)

Venter of anal legs with distinct, large, heavily
     sclerotized spine-like setae (Fig. 14).  Head
     color pattern as Fig. -28	S. piatrix (p. 57)
Venter of anal legs lacking large, distinct, heavily
     sclerotized setae  	  7
Head with three distinct stripes  (Fig. 19); central
     stripe occasionally with yellow spot in middle;
     each notum with distinct anterior and posterior
     dark band	S. bvonta Appalachian Form
Head and thorax lacking striped color pattern	
(p.  37)
.  .   8
Head and thoracic sclerites with dark brown to black
     background color; frontoclypeus with either dis-
     tinct central spot(s) or pair of diagonal spots
     on anterior portion of sclerite 	
Head and thoracic sclerites straw brown to bronze;
     pattern of frontoclypeus different than that above.
.  .   9

.  .  10
Frontoclypeus with large central yellow spot  (Fig. 23)
     occasionally with two or, three that may be  fused
     to form longitudinal row or stripe in center of
     the sclerite (Fig. 24).  ........  S. slossonae  (p. 47)
Frontoclypeus with no central spot(s) or central
     stripes; frontoclypeus with pair of diagonal
     spots on anterolateral part of sclerite  	
     (Fig. 21)	  .  S. riola  (p. 44)

Posterior half  of frontoclypeus dark brown to black
     forming triangular mark  (Fig. 25)  . . .  . S. maoleodi  (p. 50)
Posterior half  of frontoclypeus not darker than
     rest of sclerite	11
Frontoclypeus with  three indistinct  pale  spots
      (Fig. 26); one centrally  located, others ante-
      rolateral of it.  Thoracic nota with many  con-
      spicuous, thick,  short bristle-like  setae.
      Club hairs of  abdomen .less dense,and only
      slightly thicker  than regular. hairsj,; Widespread,
      common.  ...........  .. ".'-J.  .....£.  sparna  (p.  52)
Frontoclypeus |unicolored (Fig. 27).  Thoracic nota
      with bristle-like setae fewer and not very
      conspicous._  Club hairs of abdomen more dense
      and conspicuously thicker than  regular hairs.
      Rare. .  . .  .  .  . . .,'"•>"•  ........  S. Ventura  (p.  55)
                                      14

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12  (1)     Frontoclypeus with two large upturned teeth or
                 denticles on anterior margin (Fig. 35). .,,,„, „ , , 13
12'         Anterior margin of frontoclypeus straight or
                 convex; without upturned teeth. ........... f , 3,4

13  (,12)    Frontoclypeus typically with large V-shaped mark
                 (Fig. 35A); entire posterior one^fourth of
                 head yellow; lateral and ventral aspects of
                 of genae with little dark pigmentation,
                 mostly yellow (Fig. 35B). ,......,,  fff QFtfis' (p, 71)
13'         Frontoclypeus with two large anterolateral spots
                 (Fig. 36A); wide dark areas adjoining epi"
                 cranial arm; lateral, ventral, aspects of
                 genae mostly dark with only a narrow yellow
                 band behind eye (Fig. 36B)	, . . . #. &&few' (p. 75)

14  (12')   Head black with distinct carina (Fig. 33A-33B).
                 Known only from mountains of North Carolina
                 and northern Georgia	#, cgppHna (p, 68)
14'         Head variously patterned or may be black but
                 lacks distinct carina	  . ,  . , , . J,5

15  (14')   Frontoclypeus produced into a low, wide angle
                 (Fig. 38)	  fl, phalewatZ (p., 78)
15'         Anterior margin of frontoclypeus straight or,
                 at most, broadly rounded. ....... f ,.,.,,,, 3,6,

16  (15')   Large, conspicuous and heavily sclerotized stput
                 setae on venter of anal legs (Fig, 14) f ...... , . . 3,7
16'         Large and heavily sclerotized stout setae absent
                 from venter of anal legs. ............... , £2
17  (16)
17'
18  (17')
18'
Center of anterior ventral apotome produced into a
     rounded tubercle (Fig. 16) best observed in
     lateral view.  Minute spines on dorsum of ab"
     domen reaching fourth segment on which they
     are very small.  Head pattern as in Fig. 47,
     Larva extremely large.  Known only from
     central Virginia. . ........ ... H,
Anterior ventral apotome without large, rounded
     tubercle.  Minute spines reaching at least
     to segment- five.  Head pattern different
     from Fig.":47 ...... -....,.....,,
Numerous blacky7lbristle-like setae on posterior half
     of frontoclypeus (Fig. 13).  Stout spine-like
     setae on venter of anal legs more long and
     slender.  Color pattern of head as in Fig. 41  ,
     	  H,
Posterior half of frontoclypeus lacking conspicuous
     stout, bristle-like setae.  Stout spine-^like
     setae on anal legs shorter, stouter, and heavier
                                                                        (p, 98.)
                                                                           18
fff (p.  85)

   .  ,  19
                                      15

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19  (18')

19'
20  (19)


20'


21  (19')
21'


22  (16')


22'



23  (22)
23'
24  (22')
Head pattern as in Figs. 37 and 51.  All thoracic
     nota pale yellow	20
Head with more extensive brown pigment (Figs. 43,
     50).  If thoracic nota are pale yellow the
     pronotum is at least patterned and/or freckled
     with brown and darker than other thoracic nota	21

Stout, spine-like setae on venter of anal legs same
     size, shape, and color as those on ventral
     sclerites of ninth segment	 . . H. aevaba (p. 77)
These spine-like setae much smaller and less robust
     than those on venter of ninth segment . .  H. fvisoni (p. 93)

Head pattern as in Fig. 50; anterior half of fronto-
     clypeus with two rows of large, yellow spots;
     first row with three spots, second typically
     with four; posterior half of sclerite mottled
     with brown.  Known only from Harpeth River,
     Tennessee	#. patera (p. 104)
Frontoclypeus lacking two rows of pale spots
     anteriad (Fig. 43)	H. sodlca>is (p. 87)

Posterior angle of frontoclypeus with elevated mound
     or tubercle best seen in lateral or posterior
     view of head (Fig. 31B) f . . .	  . 23*
Posterior angle of frontoclyepus level with rest of
     posterior half of sclerite; no mound or
     tubercle present (Fig. 13)	 24*

Sides of head evenly curved; head typically
     unicolored dark brown to black except for
     light area around eye (Fig. 30) and occa-
     sionally behind eye.  (See diagnosis under
     H. bettewi-s p. 62)	H. betten-i (p. 61),
     	 H. depravata (p. 63), and H. potomaeensis (p. 64)
From dorsal view sides of head constricted
     centrally and widened anteriad (Fig. 31);
     posterior area of head not as wide as
     anterior; head mostly dark brown with
     pair of large, diagonal, tear-shaped spots
     (Fig. 31); sides and top of head hear epi-
     cranial stem with several dark brown, oval
     muscle scars. ......... . ,-s: :•. • • H. elissoma (p. 65)
Frontoclypeus with many stout, bristle-like setae
     conspicuous on body of sclerite; :most abun- .
     dant on posterior half of sclerite	
                                                                           25
      This mound or tubercle is often difficult to see, especially in the
black-headed bettent-type. species.  It is best observed in dorsolateral view,
and after the dorsum of the head has been cleaned of debris.
                                      16

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24'
Bristle-like setae, if present, restricted  to
     anterolateral corners of frontoclypeus;
     posterior half of sclerite may have minute,
     clear, spine-like setae but lacks larger
     bristle-like setae.  .	
                                                                           27
25  (24)
25'
26  (25')
26'
27  (24'
27'




28  (27')

28'


29  (28)
Head almost entirely dark brown to black except
     for areas around and behind eyes  (Fig. 48A);
     light areas behind and around eye forming a
     "duckling-shaped" mark (Fig. 48B); fronto-
     clypeus with pair of small, often obscure
     spots mesal to tentorial pits             H. leoncfrdi  (p.100)
Head not dark brown to black, either lighter
     brown with spots (Fig. 39), or with pattern
     shown in Fig. 45; no "duckling" mark on side
     of head; spots on frontoclypeus larger and
     more distinct	*  . 26

Frontoclypeus dark brown with two pairs of dis-
     tinct, subequal yellow spots (Fig. 39); top
     and sides of head lacking conspicuous yellow,
     oval muscle scars	H. diecoMfaa  (p. 80)
Frontoclypeus brown with a single pair of large,
     less distinct yellow spots (Fig. 45); top
     and sides of head with numerous conspicuous,
     yellow, oval muscle scars . . . . H. miss-i-ssippieHs-is  (p. 95)

Appressed thin, short, black setae sparse on all
     abdominal segments; abdomen with distinctive
     sheen-like appearance.  Entire top and sides
     of head uniform bright brownish red except
     for fine pattern of yellowish spots (Fig. 34);
     frontoclypeus with pair of indistinct central
     spots	• H. Guani-s  (p. 70)
Appressed thin, short, black setae densely covering
     all segments.  Color,,pattern unlike that of
     Fig. 34; frontaelypeus with "or without pair
     of central pale sppts
                                                               28
                           nas tuabdominal segments
Scale hairs sparse-on at
     I - IV (Fig. 11). . .\ . '.v*_^_. . .	 29
Scale hairs very ^abundant on all segments, may be
     fewer in number on segments I - II (Fig. 9-10)	31
            -...'. A "":"•..              •
Entire abdomen 'virtually lacking scale hairs, very
     few found dti last two segments.  Frontoclypeus
     with two pairs of distinct, subequal yellow
     spots (Fig. 40); one pair centrally located,
     the other pair anterolateral to these; top and
     sides of genae with many oval yellow muscle
     scars encircled with dark brown pigment	
     	H. ' dernova (p. 83)
                                      17

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29'
30  (29»
Scale hairs on at least the last three segments.
     Head pattern unlike that of Fig. 40; muscle
     scarsj if present, are solid brown. ....
                                                                           30
30'
31  (28')
31*
32   (31')
 32*
i?r6m dorsal view sides of head constricted cen-
     trally and widening anterior to this (Fig.
     32); posterior portion of head not as wide
     as anterior pottion; frontoclypeus with
     two large, tear-shaped spots; dorsolateral
     aspects of genae with numerous solid brown
     muscle scars.	H. deoalda (p. 66)
Sides of head evenly rounded; head only slightly
     wider anteriad; frontoclypeus more or less
     unicolored (Fig. 42); dorsolateral aspects
     of head iwth a few indistinct yellow muscle
     scars	•  H. ayinaie (p. 86)

Frontoclypeus with two pairs of yellow spots,
     one pair centrally located, the other pair
     anterolateral to these (Fig. 46A); spots
     often fused to form large, diagonal streaks
     on anterior portion of sclerite; posterior
     half of sclerite mottled; in lateral view
     dark area behind eye with three to four
     horizontal rows of yellow muscle scars
     curved dorsad posteriorly (Fig. 46B); dark
     pigment behind eye contiguous with dark
     pigment on venter of head	H. venularis (p. 96)
Frontoclypeus with single pair of centrally
     located spots; yellow muscle scars behind
     eyes not in rows; at least a narrow yellow
     band separating dark pigment behind eye
     and that of venter of head; posterior half
     of frontoclypeus solid dark brown, not
     mottled .	32

Dorsum of head dark brown with two small, often
     inconspicuous spots  (Fig. 49A); venter of
     head mostly dark brown, except for two
     quadrate yellow spots adjacent to gular
     suture; pigmentation of dorsum and venter
     of head separated by yellow band behind
     eyes to back margin  of head; area around
     eyes yellow	•  •  H. hageni  (p.  102)
Frontoclypeus dark brown with pair of large,
     conspicuous yellow spots; large, yellow
     areas on ventrolateral aspects of head
      (Fig. 3 and 44)	•	
      »»..„„*.  H.  sirmlans  (p. 90) and H.  inoomnoda (p. 92)
                                       18

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                  Mesonotum
                              Metanotum
          Pronotum
                                                    Pleural Gill
Fore Trochantin
                                           .,-x.-:,:•.: -^ff^'f- v^-i t:t>>
                                          s^^i?'flS
                                               •  ros'"    -.•- v


                                                 '
      Anal Leg
                       •.... ,1
                    Anal Papaliae
                                                 Ventral  Gills
Fig.  1,  Hydpopsyehe 'bet-bent larvae, lateral aspect (adapted and modified from
Ross,  1944).
                                   19

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                                                 Mandible
                                                 	-Maxillary Cardo

                                                         Submentum
                                                 Anterior  Ventral Apotome

                                                 Genae

                                                 Stridulatory Surface
                                                    .Gular Suture

                                                     Labrum
               Posterior Ventral Apotome

                          Mandibles

                   Frontoclypeus
                                             Prosternum
                                                     Poststernal Plates
Fig.  2-6, Hycfropsyche simulans-, Fig.  2,  larva, head, ventral aspect; Fig.  3,
larva,  head, dorsal aspect;  Fig. 4, larva, labrum,  dorsal aspect; Fig.  5,
larva,  mandibles,  dorsal aspect; Fig.  6, larva,  prosternum  and poststertial
plates.
                                          20

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                                                       8
Fig. 7 and 8, Hydropsyohe sp.; Fig. 7, pupa, case, dorsal aspect; Fig. 8,
pupa, case, ventral aspect.
                                      21

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Fig.  9.  Hydropsyohe simulans larva, apex of abdomen, dorsal aspect
          (modified from Ross, 1944).

Fig. 10.  Hydropsyohe simulans larva, abdominal segment VII, dorsal aspect,
          enlarged portion of epidermis (modified from Ross, 1944).

Fig. 11.  Hydropsyche opinale larva, abdominal segment VII, dorsal aspect,
          enlarged portion of epidermis (after Ross, 1944).

Fig. 12.  Symphitopsyche peeurvata larva, abdominal segment VII, dorsal
          aspect, enlarged portion of epidermis (after Ross, 1944).

Fig. 13.  Hydropsyohe d-ioantha larva, head, lateral aspect; arrow shows
          bristle-like setae on genae and frontoclypeus.

Fig. 14.  Hydropsyohe aerata larva, apex of abdomen, ventral aspect (after
          Ross, 1944)

Fig. 15.  Hydropsyohe sp. larva, frontoclypeus; A. dorsal aspect showing
          small spine-like setae on posterior portion of sclerite; B.
          lateral aspect.

Fig. 16.  Hydropsyohe. hoffmani larva, head, lateral aspect; arrow shows
          tubercle on Anterior Ventral Apotome.
                                       22

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10
                          23

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Fig. 17.  Symphitopsyche b-lf-Lda larva, head,  dorsal  aspect.

Fig. 18.  Symph-itopsyche walkeri variant  larva, head,  dorsal  aspect.

Fig. 19.  Sywphitopsyche bx>onta (Appalachian  Form) larva,  head,  dorsal aspect.

Fig. 20.  Syrnph-itopsyahe moTOsa larva, head,  dorsal  aspect.

Fig. 21.  Symphitopsyohe riola larva, head, dorsal aspect.

Fig. 22.  Symphitopsyohe alhedva larva, head,  dorsal aspect.

Fig. 23.  Symph-Ctopsyohe slossonae  larva, head, dorsal aspect.

Fig. 24.  Symphitopsyohe slossonae  variant  larva, head,  dorsal  aspect.

Fig. 25.  Symphitopsyehe maoleodi larva,  head, dorsal aspect.

Fig. 26.  Symphitopsyohe spcama. larva, head,  dorsal  aspect.

Fig. 27.  Symphitopsyche Ventura larva, head,  dorsal aspect.

Fig. 28.  SympTvitopsyohe piatrix presumed larva,  head, dorsal aspect.

Fig. 29.  Symphitopsyahe etnteri- larva, head,  dorsal aspect.

Fig. 30.  Hydropsyche betten-L larva,  head,  dorsal aspect.

Fig. 31.  Hydropsyche el-issoma larva, head; A. dorsal aspect;  B.  lateral
          aspect, arrow showing rounded tubercle  at  posterior angle of
          f rontoclypeus.                         -<••-',
                                       24

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25

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Fig. 32.  Hydropsydhe decalda presumed larva, head, dorsal aspect.

Fig. 33.  Hydropsyohe Carolina larva, head; A.  dorsal aspect; B.  dorsola-
          lateral aspect.

Fig. 34.  Hydropsyohe ouanis larva, head, dorsal aspect (after Ross, 1944).

Fig. 35.  Hydropsyohe orris larva, head; A.  dorsal aspect; B.  lateral
          aspect.

Fig. 36.  Hydropsyche bidens larva, head; A.  dorsal aspect; B.  lateral
          aspect.

Fig. 37.  Hydropsyohe aerata larva, head, dorsal aspect.

Fig. 38.  Hydropsyche phalerata larva, head, dorsal aspect.

Fig. 39.  Hydropsyohe dioantha larva, head, dorsal aspect.

Fig. 40.  Hydropsyche demora larva, head, dorsal aspect.

Fig. 41.  Hydropsyohe valanis larva, head, dorsal aspect.

Fig. 42.  Hydropsyohe arinale larva, head, dorsal aspect  (after Ross, 1944),

Fig. 43.  Hydropsyohe scalaris larva, head, dorsal aspect.
                                      26

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35A
35B
36A
36B
                      27

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Fig. 44.  Hydropsyche simuLans larva, head; A.  dorsal  aspect;  B.   lateral
          aspect.

Fig. 45.  Hydropsyahe rrriss'issippi>ensis larva, head, dorsal  aspect.

Fig. 46.  Hydropsyehe venular-Ls larva, head; A.  dorsal aspect;  B.   lateral
          aspect.

Fig. 47.  Hydropsyshe hoffmani larva, head, dorsal aspect.

Fig. 48.  Hydropsyohe leonardi larva, head; A.  dorsal  aspect;  B.   lateral
          aspect.

Fig. 49.  Hydropsyohe hageni larva, head; A.  dorsal aspect;  B.  lateral
          aspect.

Fig. 50.  Hydropsyohe pcrbeva larva, head, dorsal aspect.

Fig. 51.  Rydvopsyohe 'f-plsam, larva, head, dorsal aspect.
                                      28

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 44A
                       44B
 46A
46B
48A
                     48B
                      49B

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                                  SECTION 8

                            DISCUSSION OF SPECIES

                          The Genus Symphitopsyohe


                        Symphitopsyohe bifida (Banks)
                                  (Fig. 17)

     Symphitopsyohe bifida (Banks), 1905.  Tr. Amer. Ent. Soc., 32:15 (Type
locality:  -"Fort Collins, Colorado").

     As Hydropsyche bifida:  Banks, 1907.  Cat.  Neur. Ins. U. S., p. 47;
Ulmer, 1907.  Gen. Ins., 60:171; Essig, 1926.  Ins. W. N. Amer., p. 177;
Batten, 1934.  Stud. N. Am. Trich., 3:76 (As syn. of H. chlorotica Hagen);
Ross, 1938c.  Psyche, 45:16; Denning, 1943.  Ent. Amer., 23:110, 112, 129-
131; Ross, 1944.  111. Nat. Hist. Surv. Bui., 23:6, 87, 91, 96, 97, 294;
Leonard and Leonard 1949.  Occ. Pap. Mus. Zoo. Mich., 522:9; Morse and
Blickle, 1953.  Ent. News, 64:71; Etnier, 1965.   Ent. News, 76:146; Edwards,
1966.  J. Tenn. Acad. Sci., 41:120 (All material examined were larvae; the
identity is in question since no adults have ever been collected or reported
from Tennessee); Corbet et al., 1966.  Can. Ent., 98:1287, 1290; Longridge
and Hilsenhoff, 1973.  Wise. Acad. Sci., Arts and Letters, 61:176; Nimmo,
1966.  Can. Ent., 98:691.

Description

     Head capsule length, 1.25 to 1.40 mm; head capsule width, 1.20 to 1.35
mm.   Seven spots in checkerboard pattern on frontoclypeus.  Dark brown to
black pigmentation on margins of genae forming epicranial arms; this pigment
expanded behind eye and reaching ventrally to level of eye.  Expanded black
area behind eye with number of yellow spots.  Posterior one-fifth of dorsum
of head yellow; large area around eye and posterad of eye, ventrad of black
mark, yellow.  Venter of head with large, dark,  inverted, Y-shaped mark on
stridulatory surfaces; large, quadrate, dark brown/spot located mesally around
gular suture.  Genae dorsolaterally with numerous stout, black bristle-like
setae.  Frontoclypeus shiny and lacking such setae on main body of sclerite;
bristle-like setae of frontoclypeus present on anterolateral corners and
posterior angle of sclerite.  Labrum black, with numerous black setae;
laterally fringed by brush,of long, yellow setae.  Mandibles dark brown.
     ^Measurements given to facilitate recognition of last or second-to-last
instar larvae»             ;
                                      30

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Nota brown; pronotum slightly darker than others.  Numerous stout, bristle-
like setae present on all three nota..  Prosternum yellow; anterior and pos-
terior margins black; prosternal and mesosternal plates broxm and moderately
sclerotized.  Legs brown; distal and proximal margins of first three segments
black.  All leg segments with numerous stout setae and longer black setae.
Abdomen brown; without minute spines and scale hairs on dorsum; numerous club
hairs present on dorsum of all abdominal segments and only slightly thicker
than normal black hairs present.  Pleural gills covered with numerous short,
black hairs.  Minute spines present on venter of last two abdominal segments
and anal legs.  Anal legs also with larger, slender, sclerotized stout setae;
setae smaller, more slender, otherwise similar to stout setae on sclerotized
patches of venter of segments VIII - IX.

Variation

     A certain degree of variation has been observed in the larvae of S.
T?if-ldat and this variation relates to the number of spots on the frontocly-
peus.  The central five spots on the frontoclypeus are consistently present,
but the medial anterior and posterior spots are variable.  Some individuals
of the same population may possess all seven spots, while others have only
six and lack either the anterior or posterior.  While still other individuals
may posses only the central five and lack both the anterior and posterior
spot.

Diagnosis

     S. bif-ida is still indistinguishable from other species which have the
checkerboard color pattern.  No characters have yet been discovered to con-
sistently separate S. ~bi,f-ida3 S. cheitonis3 S. watkeri-j S. YecT£t?vaba.3 and
the Central Form of S. bronta.

Material Examined

Illinois:
     Fox R. at Algonquin, McHenry Co., II., l-VII-76.  D. A. Etnier (several
     mmts. and larvae); S. Fork Rock R. at U. S. 51, S.,of Rockford,
     Winnebago Co., II., 25-VIII-76.  D. A. Etnier (4 mmts. and several
     larvae).

Minnesota:
     Stoney R., at Mn. 1, Lake Co., Mn., 16-VII-75.  D. A. Etnier (5 adult
     males, 5 mmt. males); Crow R., Watertown,.Carver Co., Mn., 17-VI-75.
     D. A. Etnier (1 adult male, several mmts., many larvae).  Crow R.,
     Watertown, Carver Co.,:Mn., 27-VIII-75.  D. A. Etnier (several mmts.,
     and larvae); Mississippi, R., at Fort Snelling, Dakota Co., Mn., 19-VII-
     75.  D. A. Etnier (1 male mmt.); Saganaga Falls, Lake Saganaga, Cook
     Co.,, Mn., 3-VIII-75.  D. A. Etnier (2 mmt. males and several larvae);
     Sunrise R.,, at Mn. 95, Chisago, Chisago, Mn., 6-VII-76.  D. A. Etnier
     (several mmts. and larvae); Chippewa R., at Mn. 15, Montevideo,
     Chippewa Co., Mn., 8-VII-76.  D. A. Etnier (several mmts. and larvae).
                                      31

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Wisconsin:
     Eau Claire R., 2 mi. above Chippewa R., Eau Claire, Eau Claire Co., Wi.,
     28-VIII-75.  D. A. Etnier (2 .mmt. males); Eau Claire R., at Co. Rd. K.,
     Eau Claire Co., Wi., 3-VII-76.  D. A. Etnier (1 mint, male, 7, larvae). ,

Ontario, Canada:
     Northern Lights Rapids, Lake Saganaga, Ontario, Canada, 15-VIII-76.
     D. A. Etnier (6 rants.)

Distribution

     The distribution records of S. bifida indicate a northern range across
most of North America.  Published records include:  British Columbia, Colo—,
rado, Illinois, Minnesota, New Hampshire, New York, Oklahoma, Ontario, Ten-
nessee, Washington, and Wyoming.

     Edwards (1966) reports this species from Tennessee; however, this record
is regarded as dubious since no adults and only larvae were reported.  S.
bifida appears to be replaced in the midwest and southeast by S. ehe-ltonls
which inhabits rivers and streams very similar to that of S. bifida.  In all
probability, it was S. cheilonis that Edwards collected, rather than S.
bifida larvae..

Biology

     Symphitopsyche bifida is commonly collected in northern medium-sized
streams to small rivers.  These streams are typically of medium gradient with
coarse gravel to small rock substrate, rich in suspended organic materials.

     Pupal cases cylindrical, in shape made of coarse sand grains and small
rocks.  This species apparently is univoltine and Ross (1944) indicates that
emergence takes place from May to August.

Remarks

     Corbet et al.  (1966) state:  "In our opinion (S. bifida, S. bronta, and
S. morosa) these three are so closely related that their status as distinct
species is uncertain.  Many specimens agree with Ross* drawings, but some
appear intermediate.  Although we recognized three categories during this
work, we consider the taxonomy of these species requires critical
investigation."

     During the course of this work, specimens of all three species have been
examined, and it is concluded that these three species are distinct.  This
has been supported by.the larvae of the individual species involved.  5.
movosa is the most distinct of the checkerboard-patterned species on the
basis of three broken spots at the posterior angle of the frontoclypeus.
Metamorphotypes and larvae have been examined from a large portion of the
known range of the species, and the posterior three spots have never been
observed to vary.  In several localities in Minnesota where S. morosa and S.
bifida larvae and mature pupae have been taken sympatrically, no intermediate
forms have been found in either the larvae or developing male genitalia.  In

                                      32

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the Little River in Tennessee and a number of other localities, S. moicosa and
the Appalachian Form of S. bvonta are sympatric.  Here, too, intermediate
forms of either the larvae or adult genitalia have never been found.  This is
not to say that hybridization does not take place between these species and
that intermediate forms do not exist, but it is to say that, on the basis of
this work, we believe that these three species are valid.


                       Symphitopsyche eheilon-is (Ross)

     Symph-ltopsyche che-llan-ls (Ross), 1938a.  111. Nat. Hist. Surv. Bui., 21:
149-150 (Type locality:  along Salt Fork R., Oakwood, II., 18-VII-33, Ross
and Mohr).

     As Hydropsyche ehe-i1on-is\  Ross, 1944.  111. Nat. Hist. Surv. Bui., 23:
10, 87, 91, 96, 98-99, 294; Leonard and Leonard, 1949.  Occ. Pap. Mus. Zoo.,
Mich., 522:9; Etnier, 1973.  J. Ga. Ent. Soc., 8:273; Longridge and Hilsenhoff,
1973.  Wise. Acad. Sci., Arts and Letters, 61:176; Resh, 1975.  Trans. Ky.
Acad. Sci., 36:12; Tarter, 1976.  Limn. W. Va., p. 130.

Variation

     Unlike S. b~Lfi-dat which shows some variation in the number of spots on
the frontoclypeus, S. chei-lonis has never been observed to vary in this regard.
With respect to other characters, no-variation has been found in this species.

Diagnosis

     Presently, S. ohe-ilonis larvae cannot be distinguished from the larvae of
S. bif-ida, S. waLkern-, S. vecu-rvata, and the Central Form of S. bz>onta (see
Fig. 17).

Material Examined

Ohio:
     Greater Miami R., at Island Pk., Dayton, Montgomery Co., Oh., 6-VII-76.
     G. A. Schuster (several mmts., many larvae); Stillwater R., Deweese Pk.,
     Dayton, Montgomery Co., Oh., 6"-VII-76.  G. A. Schuster (several mmts.,
     many larvae); Greater Miami R., at Island Pk., Dayton, Montgomery Co.,
     Oh., 7-VIII-76.  G. A. Schuster, F. Schuster (several mmts., many larvae).

Tennessee:
     Bull Run Cr., on Tn. 162, just S. of Oak Ridge, Knox Co., Tn., l-IV-76.
     G. A. Schuster, D. A. Etnier (2 male mmts., many larvae); Bull Run Cr.,
     on Tn. 162, just S. of Oak Ridge, Knox Co., Tn., 18-IV-76.  D. A. Etnier
     (several mmts. and larvae).

Virginia:
     Craig Cr.,'on Va. 615, Craig Co., Va.j 28-V-75.  G. A. Schuster, D. A.
     Etnier (2 male mmts., 1 female mmt., many larvae).
                                      33

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Distribution

     Published records of S. ehei,1onis include the following states:  Illinois,
Indiana, Kentucky, Michigan, Tennessee, and West Virginia.  The above new dis-
tribution records from Ohio and Virginia should be added to this list.

Biology

     S. chei/lonis appears to be quite tolerant of high levels of suspended
organic materials, for the streams from which it has been collected have high
concentrations of these substances.  The streams in which S. ahei.lori'is has
been taken have slow to moderate currents with large riffle areas.  They are
typically warm water streams which do not normally contain Symphitopsyehe
species.  Symphttopsyehe consists primarily of species which seem to prefer
cold water to spring-like situations.  S. che-ilonis is one of the few species
in this group which lives regularly in warm water streams.

     The emergence period for S. oheildnis is from early April through
September,


                      Symphitopsyahe vecuwata (Banks)

     Symphitopsyehe recuwaba (Banks), 1914.  Can. Ent., 46:253 (H. slossonae
var.)   (Type locality:  "Go Home Bay,. Ont., Split Rock, June 9, E. M. Walker").

     As Hydropsyehe recurvatai  Sibley, 1926.  Bui. Lloyd Libr., 27 Ent. Ser.,
5:104; Betten, 1926.  Mem. Cornell Ag. Exp. Stat., 101:524; Betten, 1934.
Caddisflies N.Y. St., p. 190; Neave, 1934.  Int. Rev. Hydrobiol., 31:159, 161,
169; Milne, 1936.  Stud. N. Am. Trich., 3:73 (as syn. of slossonae); Ross,
1938.  Psyche, 45:18.  (descript. of type); Marshall, 1939.  Ann. Ent. Soc.
Am., 32:668, 671, 67a; Ross, 1941.  Trans. Am. Ent. Soc., 67:93; Denning,
1943.  Ent. Amer., 23:110, 112, 126-127; Milne, 1943.  Can. Ent., 75:192;
Ross, 1944.  111. Nat. Hist. Surv. Bui., 23:10, 14, 87, 91, 96, 99, 294;
Leonard and Leonard, 1949.  Occ. Pap. Mus. Zoo., Mich., 522:10; Etnier, 1965.
Ent. News, 76:146; Nimmo, 1966.  Can. Ent., 98:691; Corbet et al., 1966.
Can. Ent., 98:1291; Blickle and Morse, 1966.  Me. Ag. Exp. Stat. Bui.,24:6;
Longridge and Hilsenhoff,, 1973.  Wise. Acad. Sci., Arts and Letters, 61:176.

     As Eydropsyohe codona:  Betten, 1934.  Caddisflies N.Y. St., p. 187;
Milne,  1936.  Stud. N. Am. Trich., 3:73  (as syn. of slossonae); Ross, 1938.
Psyche, 45:18 (as syn. of reourvata).

Variation                                       ,c,

     The variation found in this species is similar to the variation reported
by Ross (1944).  He states:  "Head varying from the dark checkered pattern to
almost  entirely yellow with a few brown markings outlining a skeleton
checkerboard . . . ."
                                      34

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 Diagnosis

      This widespread species can not be separated from S. bifida, S. oheilonis,
 S.,Walkepi, and the Central Form of S. brcnta on the basis of present know-
 ledge of Symphitopsyche larval taxonomy (see Fig. 17).

 Material Examined

 Minnesota:
      Stoney R., at Mn 1, Lake Co., Mn., 16-VIII-75.  D. A. Etnier (5 mmts.
      males, 1 adult male); Mississippi R., at Fort Snelling, Dakota Co., Mn.,
      19-VII-75.  D. A. Etnier (1 male mint.); Saganaga Falls, Lake Saganaga,
      Cook Co., Mn., 3-VIII-75.  D. A. Etnier (3 male mmts.); Saganaga Falls,
      Lake Saganaga, Cook Co., Mn., 2-VIII-75.  D. A. Etnier (5 male mmts.,
      several larvae); Sunrise R., at Mn. 95, Chisago Co., Mn., 6-VII-75.  D.
      A. Etnier (1 male mint., several larvae).
 Ontario:
      Northern Light Rapids, Lake Saganaga, Ontario, Canada, 15-VIII-76.
      A. Etnier (6 mmts.).
D.
 Wisconsin:
      Eau Claire R.,  2.0 mi. above Chippewa R., Eau Claire, Eau Claire Co.,
      Wi., 28-VIII-75.  D. A. Etnier (1 adult male); Eau Claire R., at Big
      Falls, Eau Claire Co., Wi., 3-VII-76.  D. A. Etnier (4 male mmts.,
      several larvae); Red Cedar R. at Co. Rds. M and W, 22 Mile Ford, Dunn
      Co., Wi., 3-VII-76.  D. A. Etnier (7 mmts., numerous larvae).

 Distribution

      This widespread species is known from:  Illinois, Manitoba, Michigan,
 Minnesota, New York, Ohio, Ontario, Quebec, Saskatchewan and Wisconsin.

 Biology

      This species is widespread and has often been recorded in the literature,
 yet little is known of its biology.  As with all Symph-itopsyche species, 5".
 reewvata is found in riffle areas of rivers, but Ross (1944) noted that this
 species lives in the wave-washed areas of the Great Lakes as well.  Ross also
 indicates that S. Teourvata is more commonly collected in fast, cold rivers.

      Ross (1944) reports this species as emerging from May to September.


                 Symphitopsyehe waTkevi (Betten and Mosely)
                             ,   ._   (Fig. 18)

     Symph-itopsyohe walkeri  (Betten and Mosely), 1940.' Walker Types Trich.
Br. Mus., p. 23-25 (Nov. Norn for macu1-Lcoz>m,s Walker)  (Type locality:  "St.
Martin's Falls, Albany R., Hudson Bay").
                                      35

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     As Hydropsyohe walkeri.:  Ross, 1944.  111. Nat. Hist. Surv. Bui., 23:87,
93, 96-97, 294; Leonard and Leonard, 1949.  Occ. Pap. Mus. Zoo., Mich., 522:
10-11; Morse and Blickle, 1953.  Ent. News, 64:71; Etnier, 1965.  Ent. News,
76:146; Blickle and Morse, 1966.  Me. Ag. Exp. Stat. Tech. Bui., 24:6; Nimmo,
1966.  Can. Ent., 98:691; Corbet et al., 1966.  Can. Ent., 98:1288, 1291;
Longridge and Hilsenhoff, 1973.  Wise. Acad. Sci., Arts and Letters, 61:176.

     As H. maculioorn-is Walker:  Walker, 1852.  Cat. Br. Mus. Neur., 1:113;
Hagen, 1861.  Syn., Neur^ N. Am., p. 289; MacLachlan, 1863.  Ent. Ann., p. 163;
Hagen, 1864.  Verh. Zoo. Bot. Ges., 14:823; Banks, 1892.  Trans. Am. Ent. Soc.,
19:367; Ulmer, 1905.  J. Insbiol., 1:68; Ulmer, 1907.  Gen. Ins., 60:171;
Banks, 1907.  Cat. Neur. Ins. U.S., p. 47; Betten, 1937.  Caddis Flies N.Y.
St., p. 188; Milne, 1936.  Stud. N. Am. Trich., 3:73 (as syn. of alteTnans').

     As Species One:  Betten, 1934.  Caddis Flies N.Y. St., p. 192;  Betten
and Mosely, 1940.  Walker Types Trich. Br. Mus., p. 23 (as syn. of walkeri).

Variation

     Specimens of S. waTkevi, collected in Virginia show no variation in the
head capsule color pattern.  However, specimens collected from Minnesota,
especially the population from the Temperence River, possess a striking amount
of variation.  Individuals from this population range from the typical checker-
board color pattern to the pattern depicted in Fig. 18.  In this population it
appears that the latter color pattern is most prevalent.  Collections of S.
waVkevi from the Baptism River site do not show this large degree of variation.
Specimens from the Baptism River show the more typical checkerboard pattern
and cannot be separated from the other larvae with that pattern.

Diagnosis

     Symphitopsyche waikevl larvae cannot be discerned from the larvae of S..
bif-ida, S. recurvata, S. eheilonis, and the Central Form of S. bronta (see
Fig. 17).                ;
                         |
Material Examined

Minnesota:
     Temperence R., on U.S. 61, Cook Co., Mn., 21-VII-75.  D. A. Etnier (5
     male mmts., several larvae); Baptism R., at Finland, Lake Co., Mn., 9-
     VII-76.  D.. A. Etnier, M. A. Etnier, S. A. Etnier (1 male mmt., several
     larvae).

Virginia:                :                       .,-.
     Roanoke R., at Dixie Caverns on U.S. 11, lQ-:Jmi> W. of Salem, Roanoke Co.,
     Va., 28-V-75.  G. A. Schuster, D. A. Etnier  (1 male mmt.); Roanoke R.,
     at Va. 419, in Salem, Roanoke Co., Va., 5-VII-75.  D. A. Etnier (several
     larvae); Roanoke R., at Va. 419 in Salem, Roanoke Co., Va., 10-VII-76.
     G. A. Schuster (several male and female mmts., many larvae).

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Distribution

     5. walkepi. has been reported from New York, Ontario, Wisconsin, and
Quebec.  The above Virginia collections represent a substantial increase in
the range of this species.

Biology

     The northern localities from which S. walkevl has been collected repre-
sent medium sized to small rivers.  These streams typically have a medium
gradient with coarse gravel to small rock substrate and are rich in suspended
organic materials.  Both Minnesota larval localities are trout streams.  The
Roanoke River locality in Virginia is very similar to the habitat described
above.  The locality from which most of the material has been collected in
Salem, on Va. Hwy. 419, has a large riffle area in which S. walkeri, is common.
In the same riffle S. bronta is also present but not in the abundance of S.
walkeTi-,  Downstream from this riffle is a long, smooth run in which there
are many large rocks on which Hydropsyche leonardi and H. hoffmani are common.
Symphitopsyohe walkeri was not collected in this area, suggesting that this
species may prefer small rocks in the faster, more oxygenated riffle areas.

     Published records indicate that this species emerges at least from May
to August.
                         SympM-topsyche bronta (Ross)
                                  (Fig. 19)

     SympTrltopsyche bronta (Ross), 1938a.  111. Nat. Hist. Surv. Bui., 21:
149 (Type locality:  Branson, Mi., 19-V-36, along Prairie R., Prison and
Ross).

     As Hydropsyche bronta: • Denning, 1943.  Ent. Amer., 23:110, 112, 125-126;
Ross, 1944.  111. Nat. Hist. Surv. Bui., 23:6, 9, 87, 91, 95, 98, 294; Leonard
and Leonard, 1949.  Occ. Pap. Mus. Zoo., Mich., 522:9; Etnier, 1965.  Ent.
News, 76:146; Blickle and Morse, 1966.  Me. Ag. Exp. Sta. Bui., 24:6; Corbet
et al., 1966.  Can. Ent., 98:1287, 1290; Nimmo, 1966.  Can. Ent., 98:691;
Unzicker et al., 1970.  J. Ga. Ent. Soc.., 5:171; Longridge and Hilsenhoff,
1973.  Wise. Acad. Sci., Arts and Letters, 61:176; Resh, 1975.  Tr. Ky.
Acad. Sci., 36:12.

     The species known as Symph-itopsyohe bronta. presents a unique problem to
the larval taxonomy of the genus.  Two distinct larval forms have been col-
lected and associated to the adult of S. bronta.  The genitalia of both forms
have been critically compared by Dr. John Unzicker with the holotype of S.
bronta, and found to be identical.  The two forms, one of which shall be
called the Central Form and the other the Appalachian Form, are distinct on
the larval level on the basis of coloration.  The Central Form, which has
been collected and associated from Ohio, Indiana, and Illinois, possesses
the typical checkerboard pattern and cannot at this time be differentiated
from S. b-i-f-Cda, S. walker-C, S. recurvata, or S. dheilonis.  The Appalachian
Form has been collected, associated, and reported from the Appalachian system,

                                      37

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from New Hampshire  to Tennessee  and North. Carolina.  The Appalachian Form is
a striking  larva with a  zebra-striped  color pattern  that can readily be dif-
ferentiated from the Central Form.  Adult genitalia  of these two forms have
been extensively examined and  compared, and no  characters have yet been ob-
served  to separate  them  consistently.  The larvae are distinct and intermedi-
ates of the two forms have never been  collected.  It is quite possible that
this situation represents two  sibling  or  cryptic species which cannot be
separated at the adult level but are easily discriminated in the larval stage.
It is unfortunate that insufficient material has been available for study to
state with  certainty whether this is the  case or not.  The situation requires
further study, and  a great amount of material must be examined before a de-
finitive statement  is possible.  For this reason, the two forms are treated
as such rather than distinct species or even subspecies.

     It is  our feeling that the  critical  study  areas include Kentucky, New
York, Ohio,  and Pennsylvania—states in which an overlap of these two forms
may occur.   Furthermore, an attempt to determine the distribution of each
form on the basis of published records, for which identification was based on
adult genitalia, is very difficult.  For  example, Resh's (1975) records from
Kentucky cannot be  termed either the Central or Appalachian Form since
Kentucky lies in a very critical area  of  possible overlap of these two forms.
Except  for  Resh (1975) and Blickle and Morse (1966), all the published records
are from areas in which the Central Form  is expected.  Blickle and Morse
(1966)  present the  only published record  of what may possibly be the
Appalachian Form.
                      a.  The Central Form of S. bronta
Variation
     Only a limited number of these larvae were available for study, and,
therefore, very little can be concluded regarding variation in this form.
However, Ross (1944) states:  "In coloration similar to b-iflda, especially
with reference to the checkered type of pattern on the head.  As with bifida,
there is considerable variation in the details of this pattern."

Diagnosis

     Because the larva of this form is indistinguishable from S. bi-f-Cda, S.
walkeTi, S. Yecwcvaba, and S. ohe-llon-ls, the reader is referred to the des-
cription of the larva of S. bifida, page 30 (see Fig. 16).

Material Examined

Illinois:
     Quiver Cr., Havana, Mason Co., II., 29-V-36.  C. 0. Mohr (1 male and 1
     female mmt.); Quiver Cr., Havana, Mason Co., II., 29-V-36.  Mohr and
     Burks (1 male mmt. and 4 larvae); White Pine Pk., Ogle Co., II., 30-V-36.
     H. H. Ross (4 female mmts., 7 larvae); Apple R., Canyon St. Pk., Jo
     Daviess Co., II., 2-III-38.  Ross and Mohr (7 larvae).
                                     38

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Distribution.

      Because  so  few metamorphotypes  and associated  larvae are available, it
is extremely difficult  to present an  accurate range  for the Central Form of S.
bvonta.  Personal communications with a number of workers  (E. McElravy, Ohio;
S. Newhouse, Indiana; V. Resh, Indiana) indicate that they have collected only
the  checkerboard  form of S. bronta in their areas.   Also, the junior author,
who  has undertaken extensive study in both Minnesota and Wisconsin, has never
encountered striped Symphitopsyohe larvae from either of these two states from
which S. bronta has been reported.  It is our contention that the Central Form
is widespread  throughout the midwestern states.  However, a great deal of addi-
tional associated larval material is  necessary to substantiate this statement.

Biology

      Ross (1944) makes these comments concerning the Illinois collections of
S. bvontai  trln Illinois this species is restricted with few exceptions to
small medium sized streams in northern Illinois.  Most of these are spring fed;
all  are permanent.  The adults emerge from April to  the latter part of August."
                    b.  The Appalachian Form of S. bronta
Variation
      The only variation that has been observed in this form is the presence
or absence of a central spot in the intermediate band on the head.  The degree
of distinctiveness of the dark banding .is also slightly variable, and the
bands may fade to a certain extent after being stored in alcohol for a long
period of time.  This variation is intra-populational, and has been found in
most populations studied.

Diagnosis

      The Appalachian Form of S. bronta differs from the Central Form only by
its consistent striped head pattern.  No other characters have been found to
distinguish this form of S. bronta from the Central Form or from the other
checkerboard-patterned species.  Therefore, the description of the larva of
S. bifida (page 30) should be referred to for comments regarding characters
other than the color pattern.

      The unique color pattern of the Appalachian Form consists of distinct
stripes on the head and nota (Fig. 19).  There are typically anterior, inter-
mediate and posterior dark bands on the head.  The medial band on occasion has
a central yellow spot.  Each notum also possesses distinct dark bands on the
anterior and posterior margins.                                             '-••

      The background color for the head and thoracic sclerites is straw yellow,
contributing to the distinctiveness of the stripes.  No specimens of the
Appalachian Form have ever been collected which have a pattern approaching
that of the checkerboard pattern.
                                      39

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Material Examined

Maryland:
     Flint Stone, Allegany Co., Md., 19-IV-38.  H. H. Ross (1 male mmt.).

North Carolina:
     Shoal Cr., on U.S. 441, 1.0 mi. S. of Cherokee, Jackson Co., N.C., 1-V-
     76.  G. A. Schuster (6 larvae); Oconaluftee R., at Cherokee, Swain Co.,
     N.C., 27-VI-76.  D. A. Etnier (1 larva).

Tennessee:
     Little R., 2.0 mi. above Townsend, Blount Co., In., on Tn. 73, 14-VIII-
     73.  M. Hughes, W. Dickinson, G. A. Schuster (several larvae); Little
     Pigeon R., 1.0 mi. S. of Richardson Cove on Co. Rd. 2421, Sevier Co.,
     Tn., 21-VI-75.  G. A. Schuster (6 larvae); Little R,,, at Townsend, Blount
     Co., Tn., on Tn. 73, 19-VI-75.  M. Hughes, W. Dickinson, G. A. Schuster
     (2 male mmts., 12 larvae); Cox Property, East Slope of Webb Mtn., Sevier
     Co., Tn., 24-1-75.  R. Smith, M. ₯arren, K. Cottrell (3 larvae); W. Prong
     Little Pigeon R., above Gatlinburg, Sevier Co., Tn., in Great Smoky Mtn.
     Nat. Pk., 17-1-76.  L. and W. Starnes (4 larvae).

Virginia:
     Roanoke R., at Wayside Park, 10 mi. above Salem on U.S. 11, Roanoke Co.,
     Va., 5-VII-75.  D. A. Etnier (1 male mmt., 7 larvae); Roanoke R., at Va.
     419 in Salem, Roanoke Co., Va., 5-VII-75.  D. A. Etnier (several larvae);
     North Fork Holston R., at Saltville, Va., below sludge ponds, VII-75.
     TVA (5 larvae); Roanoke R., at Va. 419 in Salem, Roanoke Co., Va., 10-
     VII-76.  G. A. Schuster (several mmts. and larvae).

Distribution

     The exact range of the Appalachian Form of S. bronta is unknown, but it
is believed to be restricted to drainages originating in the mountain system
from New Hampshire to the Carolinas and Tennessee.  Dr. Oliver Flint (U.S.N.
M.) has informed us that he has collected this larva as far north as New
Hampshire, and Mr. Justin Frost (Rowe, Mass.) has indicated that he has col-
lected this larva on a number of occasions in the Deerfield River drainage
system in Massachusetts.

Biology                  :

     These striped larvae have been collected in a variety of habitats in
North Carolina, Tennessee, and Virginia.  The above collections from the
Little River and West Prong of the Little Pigeon River in East Tennessee re-
present localities which can be considered cold mountain streams—brown and
rainbow trout waters.  Contrasting with this, Shoal Creek in North Carolina,
from which larvae have also been collected, is a very small creek  ( 1 meter
wide) with a heavy suspended organic load and warm water in the summer
months.

     The emergence period for this form of S. DTonta appears to be from April
to August.

                         :             40

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                        Symphitopsyohe movosa (Hagen)
                                  (Fig-20)

     Symph-itopsyche movosa (Hagen), 1861.  Syn. Neur. N. Am., p. 287 (Type
localities:  "St. Lawrence R., Can.; Washington (Osten Sacken); N. Red R.
(Kennicott); Trenton Falls, N.Y. (Osten Sacken)").

     As Hydropsyohe morosai  1873.  P. Boston Soc., 15:297; Ulmer, 1907.
Gen. Ins., 60:171 (as syn. of alternans); Milne, 1936.  Stud. N. Am. Trich.,
3:71, 73; Ross, 1938.  Psyche, 45:16; Brimley, 1938.  Ins. N. Carolina,
p. 252; Betten and Mosely, 1940.  Walker Types Trich. Br. Mus., p. 21-31;
Denning, 1943.  Ent. Amer., 23:110, 112, 127-129; Ross, 1944.  111. Nat.
Hist. Surv. Bui., 23:91, 96, 98, 294; Leonard and Leonard, 1949.  Occ. Pap.
Mus. Zoo., Mich., 522:10; Morse and Blickle, 1953.  Ent. News, 64:71;- Etnier,
1965.  Ent. News, 76:146; Nimmo, 1966.  Can. Ent., 98:691; Corbet et al.,
1966.  Can. Ent., 98:1287, 1290; Blickle and Morse, 1966.  Me. Ag. Exp. St.
Tech. Bui., 24, p. 6; Longridge and Hilsenhoff, 1973.  Wise. Acad. Sci.,
Arts and Letters, 522:176; Resh, 1975.  Trans. Ky. Acad. Sci., 36:12; Tarter,
1976.  Limn. W. Va., p. 130.

     As Hytfropsyche chlorot-iea Hagen:  Hagen, 1861.  Syn. Neur. N. Am., p.
290; Hagen, 1864.  Verh. Zool. Bot. Ges., 14:821; Banks, 1892.  Trans. Am.
Ent. Soc., 19:367; Banks, 1907.  Cat. Neur. Ins. U.S., p. 47; Ulmer, 1907.
Cat. Coll. Seys 6, 1:64-65; Ulmer, 1907.  Gen. Ins., 60:171; Banks, 1908.
Proc. Ent. Soc. Wash., 9:155; Betten, 1926.  Mem. Cornell Ag. Exp. Stat.,
101:524; Betten, 1934.  Caddis Flies N.Y. St., p. 7, 186-187; Neave, 1934.
Int. Rev. Hydrobiol., 31:169; Milne, 1936.  Stud. N. Am. Trich., 3:70,  72,
723; Brimley, 1938.  Ins. N. Carolina, p. 251; Ross, 1938.  Psyche 45:16
(as syn. of movosa).

Description

     Head capsule length, 1.30 to 14.0 mm; width, 1.10 to 1.25 mm.  Fronto-
clypeus with checkerboard color pattern  (Fig. 20); posterior corner of
frontoclypeus always with a cluster of three smaller yellow spots.  Posterior
one-fourth of head yellow; remainder of dorsal surface of head with dark
brown to black background color.  Ventral surface of head mostly yellow with
wide, dark, inverted Y-shaped marks below each eye, and black bordering the
ventral gular sutures.  Genae covered dorsally and laterally with stout black
setae to posterior corner of frontoclypeus.  Frontoclypeus devoid of such
setae except for anterolateral corners.  Labrum dark brown to black, covered
dorsally by black setae with lateral fringe of straw colored hairs.  Nota
covered with short, black setae; as dark as head, except for yellowish  spot  
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 Variation

      Like several other Symphitopsyohe species with the checkerboard color
 pattern,  S.  morosa exhibits considerable variation of this pattern.   All the
 variations we have observed in this species have been in larvae collected in
 Minnesota or Wisconsin.  Variations observed in northern material  are:   (1)  a
 much lighter colored head and nota, (2)  the venter of the head  lacks dark pig-
 mentation, (3)  the anterior three or four light spots on the  frontoclypeus are
 often fused  to  form large yellow areas on that sclerite.

      Even though considerable variation exists in some populations,  we have
 always been  able to identify S.  morosa on the basis of the three small spots
 on  the posterior corner of the frontoclypeus.   We have examined a  considerable
 number of S.  morosa larvae from many populations and have found the  only vari-
 ation in  this character is that,  on rare occasion,  there may  be only two small
 spots rather than three.   Otherwise,  this character has proven  to  be reliable
 in  the identification of the H.  morosa larvae.

 Diagnosis

      At first glance, this species  fits  the checkerboard  color  pattern attri-
 buted to  several  Symph-itopsyche  species.   However,  S. morosa  can be  consis-
 tently distinguished  by the presence  of  the three small spots clustered  in
 the posterior corner  of the frontoclypeus  (Fig.  20).  No  other  checkerboard-
 patterned Sympkitopsyohe  species  exhibits  these spots.   In all  of  the other
 species,  if  any marking is present  in the  posterior corner of the  frontocly-
 peus,  it  is  a single, large,  yellow spot  (Fig.  16).

 Material  Examined

 Minnesota:
      Temperence R., at  U.S.  61, Cook  Co., Mn.,  21-VII-75.  D.  A. Etnier
      (several mmts. and larvae); Mississippi R., at Fort Snelling,  Dakota Co.,
     Mn., 19-VII-75.  D. A. Etnier  (1 male mint.); Temperence R., at U.S.  61,
     Cook Co., Mn., 31-VII-75.  D. A. Etnier  (several mmts. and larvae);
     Devil's Track R.,  at  U.S. 61, Cook Co., Mn., 13-VIII-75.   D. A.  Etnier
      (many larvae); Sunrise R., at Mn. 95, Chisago Co., Mn., 6-VII-76.  D. A.
     Etnier  (1 male mmt.).

North Carolina:
     Panther Dr., at Fines Cr. exit, Haywood Co., N.C., 24-V-75.  D.  A.  Etnier
     and G. A. Schuster  (2 male mmt.); Shoal Cr., on U.S. 441, 1 mi.  S.  of
     Cherokee, Jackson Co., N.C., l-V-76.  G. A. Schuster (2 larvae); Ocon-
     luftee R., at Cherokee, Swain Co., N.C., 27-VI-76.  D. A. Etnier (1  larva)

Tennessee:
     Little R., 4-5 mi.  N. of Townsend, Blount Co., Tn., on Tn.  73, 19-VII-74.
     M. Hughes, R. Smith, G. A. Schuster (numerous larvae); Little  R., off Tn.
     73 at Townsend, Blount Co., Tn., 27-VII-74.  M. Hughes, R.  Smith, G. A.
     Schuster (4 male mmts., 1 female mmt.); Little Pigeon R., off  Tn. 2421
     at Richardson Cove, Sevier Co., Tn., 10-VII-74.  C. and G.  A.  Schuster
     (many larvae); Little R., at U.S. 411 bridge, Blount Co., Tn., 24-VI-74.

                                    ,  42

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     C. and G. A. Schuster  (1 male mint.); Little R., 1.5 mi. S. of jet. River
     Rd. and U.S. 411, Bount Co., Tn., 18-IV-75.  G. A. Schuster  (1 male mmt.);
     Little R., 2 mi. above Townsend, Blount Col, Tn., 14-VI.I-75.  M. Hughes,
     W. Dickinson, G. A. Schuster (2  female mmt., many larvae); Little Pigeon
     R., 1 mi. S. of Richardson Cove  on Co. Rd. 2421, Sevier Co., Tn., 21-VI-75.
     G. A. Schuster  (7 larvae); Little R., at Townsend on Tn. 73, Blount Co.,
     Tn., 19-VT-75.  M. Hughes, ₯. Dickinson, G. A. Schuster (many larvae);
     Nolichucky R., at Solomon Island off unnamed Co. Rd., approx. 12.5 mi. N.
     of Newport, at Cocke and Green Co. line, 26-X-75.  D. A. Etnier (2 larvae).

Virginia:
     New R. below U.S. 460 bridge, Giles Co., Va.,  29-V-75.  D. A. Etnier, G.
     A. Schuster (1 male, 3 larvae); James R., at Peters Cr. on Va. 501,
     Bedford Co., Va., 30-VIII-76.  D. A. Etnier, G. A. Schuster  (3 mmts.,
     several larvae).

Wisconsin:
     Red Cedar R., at Co. Rd. M and W, 22 Mile Ford, Dunn Co., Wi., 3-VII-76.
     D. A. Etnier (2 male mmts.).

Distribution

     This species is widespread across the northeastern U.S. and southeastern
Canada.  In recent years it has also been found to be widespread in the south-
eastern U.S., roughly following the Appalachian Mountains south.  Its range
includes the following:  Maine, Manitoba, Michigan, Minnesota, New Hampshire,
New York, North Carolina, Ontario, Quebec, Tennessee, Virginia, West Virginia
and Wisconsin.

Biology

     This species is commonly collected in medium-sized rivers, those typified
as small-mouth bass streams.  We have collected S.  mor>osa from many localities
on the Little River, which has its origins high in the Great Smoky Mountains,
and drains into Fort Loudon Reservoir (Tennessee River).   It seems to be ab-
sent from the higher elevations where S.  slossonae and S.  alhedra are common.
The habitat there can be characterized as trout waters with large, smooth
boulders.   It is commonest in the middle portions of the river where there
are large riffle areas and where the substrate consists of small- to medium-
sized rocks covered with Podostemim sp.   Its pupal cases are most often
attached to strands of Podostemim where they are completely cylindrical in
shape.  Otherwise,  the cases are attached directly to the rock surfaces.
Symphitopsyche moTosa is uncommon in the lower reaches of the Little River
where the habitat is more like that of a large river with a high silt and
suspended particle load.  Other rivers where we have collected S.  morosa
follow the general description of the Little River of Tennessee and have
similar habitats.

     Collection records indicate that this species univoltine,  emerging from
April through September.
                                     43

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                       Symphitopsyohe viola (Denning)
                                   (Fig. 21)

     Symphitopsyche riola (Denning), 1942.  Can. Ent., 74:49 (Type locality:
Nine Mile Cr., Hennepin Co., Mn., 5-V-37.  D. G. Denning).

     As Hydropsyche T-iolai  Denning, 1943.  Ent. Am., 23:133-134; Ross, 1944.
111. Nat. Hist. Surv. Bui., 23:294; Ross and Spencer, 1952.  Ent. Soc. B.C.,
Proc., 48:46; Etnier, 1965.  Ent. News, 76:146; Longridge and Hilsenhoff,
1973.  Wise. Acad. Sci., Arts and Letters, 61:176.

Description

     Head capsule length, 1.35 to 1.40 mm; width, 1.15 to 1.20 mm.  Antero-
dorsal four-fifths of head dark brown; frontoclypeus with large triangular
anterolateral pale spots; often with a more inconspicuous central subtriangular
spot.  Area around eyes and directly posterior to them and posterior one-fifth
of head yellow.  Venter of head with large, brown, pigmented areas covering
stridulatory surfaces, and around geneal sutures.  Genae adorned dorsolaterally
with stout, short, black setae.  Anterolateral corner and margins of frontocly-
peus bordering genae with scattered stout black setae; rest of sclerite devoid
of such setae.  Labrum dark brown and covered with short, black setae; late-
rally margined with tuft of long, yellow setae.  Mandibles uniformly dark
brown.  Thoracic nota dark brown; pronotum with large lateral area slightly
lighter in color than rest of sclerite.  Each nota covered with short, black
setae and margined laterally in black.  Legs uniformly colored dark brown,
and abundantly covered with short, golden spine-like setae.  Abdomen beige,
covered with short, black hairs.  Club hairs present on dorsum of abdomen as
well as below the lateral line; not greatly thicker than other hairs.  Dorsal
sclerite and ventral surface of anal legs covered with short, black setae.

Variation

     The major variation observed in this species is the amount of contrast
of the central spot located on the frontoclypeus.  It is often completely
obscure while in other larvae it is as conspicuous as the anterolateral spots.
This variation must be kept in mind when keying this larva.  There is also
some variation in the degree of contrast of the anterolateral spots.  However,
this is not as great as in the central spot; the anterolateral spots are
always conspicuous.

Diagnosis

     This species is easily recognized by its dark brown coloration and the
pair of conspicuous triangular marks on the frontoclypeus.  However, some
specimens may have an additional central spot on the frontoclypeus which may
lead to some confusion between this species and S. spcama.  These two species
may be separated on the basis of:  (1) The nota and all leg and head sclerites
are much darker in S. riola than in S. sparna; the sclerites of S. sparna are
consistently golden to reddish brown in color; whereas, those of 5". vi-ola are
very dark brown—almost black.  (2) Due to the dark pigmentation of S. riola,
the spots on the frontoclypeus have much greater contrast with the background

                         :             44

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than those of S. spama which are often, obscured.  (3) The abdomen of S.
is darker than that of S. sparna as well as covered by many short, black hairs.

Material Examined

Minnesota:
     Middle Fork Whitewater R., at Mn. 74 bridge, 2 mi. S. of Elba, Winona
     Co., Mn., 24-VIII-76.  D. A. Etnier and family (numerous mmts. and many
     larvae).

Distribution

     This species is widespread across the northern sections of North America.
Published records include locales from the following states:  Minnesota and
Wisconsin and from British Columbia.

Biology

     Little can be said of the biology of S. piola since it has not often been
collected or studied.  The larvae and metamorphotypes are from the Middle Fork
of the Whitewater River in Minnesota, which is a medium-size trout stream.
The bottom consists primarily of coarse gravel and sand.

     Published adult collection data indicate that this species emerges from
April to September.

Remarks
     The genus Symph'Ltopsyc'he is presently being revised worldwide at the
adult level by Drs. J. D. Unzicker and H. H. Ross.  Dr. Unzicker, in personal
communications, and the junior author, have indicated the possibility that
H. Ti.o1a and H. afhedra are synonyms.  However, based on the distinctiveness
of the larvae of both forms, they are treated here as distinct species until
this question can be resolved.

                        Symph-itopsyche alhedica (Ross).
                                  (Fig. 22)

     Symphitopsyche alhedra (Ross), 1939.  Proc. Ent. Soc. Wash., 41:67 (Type
locality:  Black Gap, N.C., 24-IV-38, Ross and Burks).

     As Hydropsyche athedra-  Brimely, 1942.  Ins. N. Carolina, p. 15.     l:;.

Description

     Head capsule length, 1.65 to 1.80 mm; width, 1.45 to 1.60 mm.  Head coal
black except for circular areas around the eyes and large subrectangular
yellow area on posterior lateral aspects of.genae.  Genae and frontoclypeus
very shiny.  Genae with stout, black setae dorsally especially along fronto-
clypeus; some setae on lateral aspects of genae.  Frontoclypeus with such
setae in anterolateral corners and at posterior angle;  rest of frontoclypeus
devoid of these.  Labrum coal black with short, black setae; laterally

                                      45

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margined by tuft of long, yellow setae.  Mandibles dark brown to black.
Thoracic nota dark brown to black each adorned with numerous short, black
setae ^similar to those found on head.  Each nota laterally margined in black.
Prosternum mostly dark brown to black, laterally gold; poststernal plates
brown.  Dorsum of abdomen also with numerous club hairs, few of which are
located below the pleural gills.  Dorsal sclerite of anal legs covered with
short black setae.  Ventral aspects of anal legs also covered with such setae.

Diagnosis

     This species may be separated from other Symphitopsyche species on the
basis of head capsule coloration.  Only one other species, 5. etrtieri from
Grainger County, Tennessee, has similar coloration.  These two species can
be reliably separated by:  (1) The posterior dorsal one-fifth of the head
is slightly lighter in color than the rest of the head in S. etnieri, and
located in this area are numerous circular dark brown muscle scars (Fig. 29).
These muscle scars are absent in S. alhedra.  (2) The abdomen of S. alhedpa
is densely covered with black hairs and numerous_club hairs.  The abdomen of
S. etni.eri> has considerably less hairs, especially on the ventral surfaces,
and the hairs are thinner and shorter.  (3) The club hairs on the abdomen are
thicker and more conspicuous in S. alhedra.  (4) The average head length of
S. alhedPO. is .35 mm longer than S. etnievi, and the width is .22 mm wider.
In general, S. alhedra is more robust and larger than S. etnieri.

Material Examined

Tennessee:
     Middle Prong of Little R., at Tremont, Great Smoky Mtn. Nat. Pk., Blount
     Co., Tn., 18-XI-75.  W. C. Dickinson, G. A. Schuster (6 larvae); Middle
     Prong of Little R., at Tremont, Great Smoky Mtn. Nat. Pk., Blount Co.,
     Tn., 19-111-76.  G. A. Schuster (2 larvae); Hills Cr., on Tn. 73 between
     Gatlinburg and Cosby, Sevier Co., Tn., ll-IV-76.  D. A. Etnier (3 male
     and 1 female mmts.); Middle'Prong of Little R., at Tremont, Great Smoky
     Mtn. Nat. Pk., Blount Co., Tn., 7-IV-76.  W. C. Dickinson, G. W. Wolfe,
     G. A. Schuster (several larvae).

Distribution

     The only published record of this species is the description of the
holotype from Black Gap, N.C.  Listed above are two additional localities
from which either larvae or mature pupae were collected.

Biology

     Little is known of the biology of S.  alhedra since it has been rarely
collected.  However, it is commonly encountered in the Middle Prong of the
Little River at Tremont in the Great Smoky Mountains National Park.  The river
at this point is 12 to 15 meters wide and 0.2 to 1.0 meters in depth.  Shore
vegetation includes primarily rhododendron and shrubs.  The river bottom con-
sists of large- to medium-size boulders with little vegetation; current is

                                      46

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moderate  to  fast.  The water  is  clear with very little turbidity and has
fairly low temperatures year  round.   The river may be classified as  a brown
and rainbow  trout  stream  at this location.

     The  larvae  inhabit the faster, white-water areas which makes collecting
difficult.   Their  larval  retreats are often closely associated with  those of
Arctopsyche  -irrorata Banks.   Because  the boulders  are so  large and the current
so fast,  collecting S. alhedva is best  done by running one's fingers over the
surfaces  of  the  rocks while steadying oneself  in the current.

     Caddisflies found to be  sympatric  with S.  alhedra are.:   Bvao'hycen-tvus
sp-inae, B. lateralis, Lep-idostoma spp.,  Dolophilodes dlst-inotus,  Symph-ito-
psyohe bvonta, S.  spcama, S.  slossonae,  Aratopsyche irrorata,  Dipleetrona
modesta,  and Parapsyohe ap-Loal-is.

     Symph-itopsyche aThedra appears to  have one emergence per year.   Mature
larvae have been collected during the winter months with  few mature  larvae
collected after April.  Emergence occurs during April.

Remarks
     See "Remarks," S. riola, page 45.

                      Symph-i-topsyche slossonae  (Banks)
                              (Fig. 23 and 24)

     Symph-itopsyche slossonae (Banks), 1905.  Trans. Amer. Ent. Soc., 32:14
(Type locality:  "Franconia, New Hampshire (Mrs. Slosson)").

     As Hydropsyche slossonae:  Banks, 1907.  ,Cat. Neur. Ins. U.S.,  p. 47;
Ulmer, 1907.  Gen. Ins., 60:171; Banks, 1908.  Psyche, 15:61, 65; Betten,
1934.  Gaddis Flies N.Y. St., p. 185 (as syn. of altemans); Milne,  1936.
Stud. N. Am. Trich., 3:69, 72, 73; Ross, 1938.  Psyche, 45:18; Brimley, 1938.
Ins. N. Carolina, p. 252; Howell, 1939.  J. Elisha Mitch. Soc., 55:327; Ross,
1941.  Trans. Am. Ent. Soc., 67:92; Denning, 1943.  Ent. Am. 23:110, 112,
113, 131-133; Ross, 1944.  111. Nat. Hist. Surv. Bui., 23:8, 14, 87, 88, 96,
99, 294; Leonard and Leonard, 1949.  Occ. Pap. Mus. Zoo., Mich., 522:10;
Morse and Blickle, 1953.  Ent. News, 64:71; Ellis, 1962.  Occ. Pap. Mus. Zoo.,
Mich., 624:8; Etnier, 1965.  Ent. News, 76:146; Edwards, 1966.  J. Tenn. Acad.
Sci., 41:121; Blickle and Morse, 1966.  Me. Ag.-Exp. Stat. Tech. Bui., 24:6;
Unzicker et al., 1970.  J. Ga. Ent. Soc., 5:172; Longridge and Hilsenhoff,.-
1973.  Wise. Acad. Sci., Arts and Letters, 61:176.                         , '.:

Description

     Head capsule length, 1.45 to 1.60 mm; head capsule width, 1.25 to 1.35
mm.  Dorsum of head dark brown to black with one to three pale yellow spots
on frontoclypeus.  If two or three yellow spots present, they are always in
a longitudinal row.  Posterior one-third of dorsum of head yellow.  Dark
pigmentation of head ends just posterior to hind corner of frontoclypeus.
Genae covered dorsally and laterally with stout, black setae.  Except for
                                      47

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anterolateral corners, frontoclypeus devoid of such, setae.   Venter of head
mostly yellow with dark longitudinal marks below eyes, and surrounding suture
connecting right and left genae.  Thoracic sclerites dark brown to black and
liberally covered with stout black setae.  Nota with lateral borders edged in
black; pronotum with large rectangular light area laterally.  Sclerites of
legs golden in color; much lighter than nota or head.  Prosternum edged ante-
riorly in black, with large transverse dark spot posteriorly; rest of sclerite
yellow.  Poststernal plates dark brown and solid.  Labrum dark brown to black;
lateral setal fringe straw colored.  Abdomen brown; covered with black hairs,
and club hairs not located below pleural gills.  Dorsal sclerite of anal legs
covered with black setae; ventrally anal legs covered with long, slender,
brown spine-like setae.

Variation

     Symphitopsyche slossonae typically has the centrally located yellow spot;
however, there may be some variation to this both within and between popula-
tions.  First, it is cautioned that the spot may become nearly obscured and
overrun by the black pigmentation of the rest of the head.   After looking at
large numbers of S. slossonae, one becomes adept at recognizing such indivi-
dual variants.  Second, there is often variation with the number of spots on
the frontoclypeus.  The number may vary from one to three.   If two or three
spots are present, they will always be in a longitudinal row down the center
of the frontoclypeus (Fig. 24).  Occasionally, these two or three spots may
be fused to form one large rectangular spot.  We have found northern material
to be more variable in this regard than the southeastern material.

Diagnosis

     The color pattern of the frontoclypeus of S. slossonae is diagnostic for
this species.  There is no other known Symphitopsyche larva that exhibits this
type of pattern.  S. slossonae is easily and quickly recognized by the cen-
trally located yellow spot or spots on the frontoclypeus.  Except for the
posterior one-fourth of the head, which is usually hidden under the pronotum,
the remaining sections are very dark brown to charcoal black.  The thoracic
sclerites are nearly as dark as the head; the legs are much lighter in
coloration.

Material Examined

Minnesota:
     Devil's Track R., at Gunflint Trail, Cook Co., Mn., 31-VII-75.  D. A.
     Etnier (several•larvae); Stoney R., at Mn. 1, Lake Co., Mn., 16-VIII-75.
     D. A. Etnier  (many larvae); Temperence R., at U.S. 61, Cook Co., Mn.,
     21-VII-75.  D. A. .Etnier (3 larvae); Temperence R.,
     Co., Mn., 31-VII-75.  D. A. Etnier  (3 male mmts.,
     R., at Gunflint Trail, Cook Co., Mn., 4-VIII-75.
     several larvae).
  at U.S. 61, Cook
1 larva); Gunflint
D. A. Etnier (6 mmts.,
North Carolina:          ;
     Oconoluftee R., at Smokemont, Great Smoky Mtn. Nat. Pk., Swain Co., N.C.,
     29-IV-76.  G. A. Schuster  (many larvae); Oconoluftee R., 2 mi. N. of
                                      48

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      Smokemont, Great  Smoky Mtn. Nat. Pk.,  Swain Co., N.C., l-V-76.  G. A.
      Schuster  (many larvae);  Shoal Cr., on  U.S. 441, 1 mi. S. of Cherokee,
      Jackson Co., N.C., l-V-76.  G. A. Schuster (1  larva); Oconoluftee R., at
      Smokemont, Great  Smoky Mtn. Nat. Pk.,  Swain Co., N.C., 21-VI-76.  G. A.
      Schuster  (many larvae, several mmts.); Oconoluftee R., at Smokemont,
      Great Smoky Mtn.  Nat. Pk., Swain Co.,  N.C., 18-VI-76.  G. A. Schuster
      (many larvae, several mmts.); Oconoluftee R.,  in Cherokee, Swain Co.,
      N.C., 27-VI-76.   D. A. Etnier (several larvae).

Tennessee:
      West Piney R., Tn. 48, 7 mi. S. of Dickson, Dickson Co., Tn., 2-V-75.
      G. A. Schuster (several mmts. and larvae); Little R., 2 mi. above
      Townsend, on Tn.  73, Blount Co., Tn.,  14-VII-75.  M. Hughes, W.
      Dickinson, G. A.  Schuster  (1 male mmt.); Abrams Cr., Great Smoky Mtn.
      Nat. Pk., Blount  Co., Tn., 26-VII-75.  C. and  G. Schuster (many larvae,
      several female mmts.); Little Pigeon R., on Co. Rd. 2421, 1 mi. S. of
      Richardson Cove,  Sevier Co., Tn., ll-VI-75.  G. A. Schuster (1 larva);
      Little R., on Tn. 73 at Townsend, Blount Co.,  Tn., 19-VI-75.  M. Hughes,
      W. Dickinson, G.  Schuster  (4 larvae);  Unnamed  stream at Montgomery Bell
      St. Pk., on U.S.  70, Dickson Co., Tn., 2-V-75.  D. A. Etnier, G. A.
      Schuster (several larvae); W. Prong Little Pigeon R., above Gatlinburg,
      Sevier Co., Tn.,  17-1-76.  L. and W. Starnes (several larvae); Chimney's
      Campground in Great Smoky Mtn. Nat. Pk., Sevier Co., Tn., 3-9-VII-39.
      A. C. Cole (1 male, 3 females).
Virginia:
     Big Stoney Cr., 4 mi. above U.S. 460, Giles Co., Va., 29-V-75.
     Schuster (1 male and 1 female mmt. and many larvae).

Distribution
G. A.
     This species is widely spread over the northeastern sections of the U.S.
and Canada.  It appears to follow the Appalachian Mountains to the southeast,
where it is restricted to cold water habitats.  Published locale records
include:  Arkansas, Illinois, Maine, New York, Michigan, Minnesota, New
Hampshire, Newfoundland, North Carolina, Pennsylvania, Saskatchewan,
Tennessee, and Wisconsin.  The additional unpublished record given above is
from Virginia.

Biology

     Even though this species was described some 70 years ago, little is
known of its biology.  It is widespread over the northeastern sections of
this continent, and even when found further south, it always occurs in cold
water situations.  This pattern seems to be true for many of the Symph-ito-
psyehe species which appear to be better adapted to springs and trout-stream-
type waters than species of Hydropsyche.

     It is interesting to note that, in the Great Smoky Mountain National
Park, both S. slossonae and S. alhedra are often collected with Arotopsyche
•Crrorata Banks.  It is not uncommon to find a medium-to-large-size rock with
an Arctopsyche larval retreat attached and a larval retreat of either of the

                                     49

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above two species directly below it.  It would indeed be of interest to further
study the relationships between these SympfiitopsycTie species and A.  irrorata.
     Ross (1944) indicates that S. slossonae emerges from May to August.   Our
collection records indicate that this is also true for the southeastern
populations .

                       Symphitopsyohe maoleodi (Flint)
                                  (Fig. 25)

     Symphitopsyche maoleodi (Flint), 1965.  Proc. Ent. Soc. Wash., 67:169
(Type locality:  North Carolina, Blue Ridge Parkway, Crabtree Meadows Camp-
ground, 9-VI-61.  R. A. arid 0. S. Flint).

Description

     Head capsule length, ,1.15 to 1.30 mm; head capsule width, 1.00 to 1.15
mm.  Thoracic sclerites and sclerites of legs golden brown.  Head capsule
except for posterior one-half of frontoclypeus reddish-brown.  Posterior one-
half of frontoclypeus dark brown, producing the general outline of an arrow-
head.  Genae, both dorsally and laterally, covered with short, thin, golden
setae.  Entire frontoclypeus also covered with such setae with the greatest
number being concentrated at the posterior one-half of the sclerite.  Thoracic
sclerites and sclerites of legs covered with short, golden spine-like setae.
Labrum dark brown, covered with short, brown setae; laterally margined with
long, straw-colored setae.  Mesial border of femur covered with patch of black,
long setae.  Dorsum of abdomen covered with short, black setae and club hairs.
Club hairs found below pleural gills in numbers equal to short, black setae.
Abdominal gills sparsely branched; branches thick, sausage shaped.  Most
branching of abdominal gills from apical portion of main stem some branching
preapically.  Anal leg dorsal sclerite covered with black setae; ventrally
covered with thin, golden spine-like setae.

Variation

     We have found no distinct variability within populations or between
populations of S. maoleodi that we have examined.  Perhaps after this species
is better collected and studied,  some variations will become apparent.

Diagnosis

     This distinct species most closely resembles S. spavna with which it may
be confused.  Both species exhibit golden  coloration of the sclerites which
is quite atypical of Sympbitopsyche  species.  S. macleodi may be^easily
separated from S. spcama in a number of ways.  First, S. maoleodi  possesses
spine-like  setae over the entire  surface of the frontoclypeus, whereas, on S.
spcama these are restricted to  the anterolateral  corners of the frontoclypeus.
Second, club hairs are found below the lateral line in S. maoleodi.  These
hairs are present only on the dorsum of the abdomen in S. spcama.  Third, the
abdominal gills of S. maoleodi  are. not as  branched and subdivided  as in S.
spcama.  Last,  and most  easily  observed, are the  differences  in the color
pattern of  the head.  Symphitopsyche maoleodi is  very distinct  in  having the

                                       50

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 dark arrowhead-shaped area on the posterior one-half of the frontoclypeus;
 5. spama never exhibits this.  In turn, S. sparna commonly possesses three
 light spots on the frontoclypeus, and when these spots are indistinct, the
 frontoclypeus is unicolored.  This is never the case in S. macleodi.

 Material Examined
 Georgia:
      North Fork Chatahootchee R., Tate Br.
      0. S. Flint, Jr.  (USNM collection).
Campgrd., Rabun Co., Ga., 16-17-V-70.
 North Carolina:
      Oconoluftee R.,  3.4 mi.  W.  on Smokemont Rd.,  Great Smoky Mtn.  Nat.  Pk
      Swain Co.,  N.C.,  12-V-70.   0. S.  Flint, Jr.  CUSNM collection); Oconoluftee
      R.,  at Smokemont,  Great  Smoky Mtn.  Nat. Pk.,  Swain Co.,  N.C.,  14-VI-35.
      H. H.  Ross  (1 male mmt., Det. G.  A.  Schuster,  1976).

 Tennessee:

      George Cox  Property,  Webb Mtn., Sevier  Co., Tn.,  17-IV to 17-V-75.   G. A.
      Schuster  (many adults);  George Cox  Property, Webb Mtn.,  Sevier Co.,  Tn.,
      24-1-75.  R.  Smith, M. Warren, K. Cottrell  (many  larvae);  Horse Cr.,
      Unicoi Management  Area,  Greene Co.,  Tn.,  9-IV-76.   M.  Cox,  T.  Talek,  C.
      Manning,  C.  Knaught  (3 larvae); Spruce  Flats Br.,  Trib.  to  Middle Prong
      Little R.,  Great Smoky Mtn. Nat.  Pk., Blount Co.,  Tn., 16-IV-76.  W.  C.
      Dickinson (5 male  mmts., 4  larvae).

 Virginia:

      Grindstone  Rec. Area, 4.5 mi. W.  of  Troutsdale, Smyth  Co.,  Va.,  24-V-75.
      R. Hoffman  (USNM collection)  (several adults).

 Distribution

      Records indicate this species may be widely distributed  in  the southern
 Appalachians; however,  it has not been commonly encountered.  Its known range
 includes the following  states:  Georgia, North Carolina, Tennessee, and
 Virginia.

 Biology

     Little is known about this species.   The only published collection records
 are those given with the description of the species; included above are new
 records.  Field data indicate that this species is adapted to cold water situ-
ations.  It is most commonly  encountered in spring-type habitats but may alspTI
be found in trout waters such as the Oconoluftee River  of North Carolina.  The"
 lack of extensive branching of the abdominal gills may  be a further indicator "
that this species is adapted to cold water saturated with oxygen.  This species
needs to be collected and studied more extensively before more concrete
statements may be made regarding its biology.
                                     51

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     Collection records indicate that this species emerges from April
through July.

                        SympHitopsyche sparna (Ross)
                                  (Fig. 26)

     Symphitopsyehe sparna (Ross), 1938.  111. Nat. Hist. Surv. Bui., 21:150
(Type locality:  Lovells, Michigan, 22-V-36, along Au Sable R. Frison and
Ross).

     As Hydropsyohe sparna:  Howell, 1939.  J. Elisha Mitch. Soc., 55:327;
Denning, 1943.  Ent. Am., 23:110, 112, 134-136; Ross, 1944.  111. Nat. Hist.
Surv. Bui., 23:88, 96, 97, 294; Leonard and Leonard,  1949.  Occ. Pap. Mus.
Zoo., Mich., 522:10; Morse and Blickle, 1953.  Ent. News, 64:71; Ellis, 1962.
Occ. Pap. Mus. Zoo., Mich., 624:8; Etnier, 1965.  Ent. News, 76:146; Blickle
and Morse, 1966.  Me. Ag. Exp. Stat. Tech. Bui., 24:6; Longridge and Hilsen-
hoff, 1973.  Wise. Acad. Sci., Arts and Letters, 61:176; Resh, 1975.  Trans.
Ky. Acad. Sci., 36:12.

Description

     Head capsule length, 1.20 to 1.30 mm; head capsule width, 1.00 to 1.10
mm.  Head, nota and all leg segments golden in color; dorsum of head slightly
darker golden brown.  Frontoclypeus with a pair of yellow spots anterola-
terally, often with an additional single spot in center of the sclerite.
These spots often indiscernible and frontoclypeus then appears unicolored.
Labrum brown and covered with short, black setae; the lateral brush consis-
ting of long, yellow setae.   The genae and nota covered with many short,
black setae.  Frontoclypeus bears such setae only near geneal sutures and
anterolateral corners.  Mandibles yellow and edged in brown.  Venter of head
predominently yellow with some brown patches on stridulatory surface.
Prosternum with the posterior border black.  Poststernal plates dark brown
and solidly sclerotized.  Abdomen brownish in color with dorsum covered with
short, thin, black hairs and  club hairs.  Both types of hairs more dense on
abdominal segments I - III.   Black hairs also found on ventral surfaces of
abdomen.  Club hairs not found below pleural gills.  Ventral surface of anal
legs without spine-like setae, adorned with short, black setae.

Diagnosis

     This Symphitopsyohe species  is easily  identified and  separated  from
other species  of the group.   Symphitopsyohe sparna, S. macleod-i, and S.
Ventura are the only species  of this genus which typically have  all  sclerites
golden in coloration.  Symphitopsyche macleodi and S. spcama are readily
separated on the basis.of head capsule  pigmentation.  The  posterior  corner
of the frontoclypeus of S. macleodi is  darkly pigmented  forming  a dark brown
triangle at  this portion of  the head.  Symphitapsyehe spama  is never  pig-
mented in  this manner.

      The  frontoclypeus of S.  sparna commonly has three  light  spots,  one
located centrally,  the other two  anterolateral  to  the first.   These spots
are often indiscernible,  thus, giving  the frontoclypeus  a unicolored

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 appearance.  When  this  happens,  it may be confused with S. Ventura.  These
 two  species may be separated by  a number  of ways.  First, S. sparna has many
 conspicuous, thick,  short, black setae on the  thoracic nota; on the nota of
 S. Ventura, similar setae are not as numerous  nor are they as thick and con-
 spicuous as in  S.  sparna.  Second, the club hairs on the abdomen of S. Ventura
 are  much more numerous  and conspicuously  thicker than those of 5. sparna.
 Also, numerous  club  hairs are found below the  pleural gills in S. Ventura,
 while in S. sparna this is not the case.   Last, the head of S. sparna, except
 for  the typical three light spots on the  frontoclypeus, is unicolored;
 whereas, the dorsal  surface of the head of S.  Ventura is much darker than
 the  lateral surfaces, and the frontoclypeus is unicolored.  Symphitopsyche
 sparna is very  widely distributed and  is  commonly collected in all types of
 habitats, while S. Ventura has only been  collected and reported a few times
 since its description.

 Material Examined

 Alabama:
     Mossy Springs Cr., at Edward Parson's Farm, Limestone Co., Al., 22-V-76.
     G. A. Schuster  (many larvae); Tributary to Talapoosa R., 2 mi. S. of
     Woodland on Al. 48, Randolf Co., Al., 28-V-76.  D.  A. Etnier, G. A.
     Schuster (2 male mmts., several larvae).

 Kentucky:
     Silver Cr., on Ky. 876, 5 mi. W. of Richmond, Madison Co., Ky., 13-VI-76.
     G. A. Schuster  (several mmts. and larvae).
Minnesota: \
     North Branch Sunrise R., Chisago Co., Mn., 21-VII-75.  D. A. Etnier (3
     male mmts., several larvae); Cascade R., at &. S. 61, Cook Co., Mn.,
     13-VIII-75.  D. A. Etnier (1 male mmt., 7 larvae); Gunflint R., at
     Gunflint Trail, Cook Co., Mn., 4-VIII-75.  D. A. Etnier (many larvae).

North Carolina:
     Saluda R., at Valhalla, Haywood Co., N.C., 24-V-75.  D. A. Etnier, G.
     A. Schuster (1 male mmt., 1 larva); Shoal Cr., on U.S. 441, 1 mi.  S.
     of Cherokee, Jackson Co., N.C., l-V-76.  G. A. Schuster (several mmts.,
     many larvae); Pumpkin Town Cr., on U.S. 441, Jackson Co., N.C., l-V-76.
     G. A. Schuster (several larvae, 1 male mmt.); Oconoluftee R., at
     Smokemont, Great Smoky Mtn. Nat. Pk., Swain Co., N.C., l-V-76.  G. A.
     Schuster (8 larvae); Cullasaja R., 11.9 mi. above jet. N.C. 28 and U.S.
     441 and 23, near Franklin, Macon Co., N.C., 9-V-76.  D. A. Etnier  (4
     larvae); Cullasaja R., 6 mi. W. of Highlands, Macon Co., N.C., 19-VI-76.
     G. A. Schuster (many larvae); Oconoluftee R., at Smokemont, Great  Smoky
     Mtn. Nat. Pk., Swain Co., N.C., 27-VI-76.  D. A. Etnier (several larvae,
     1 male mmt.).

South Carolina:
     Lynches R., at Lanchester-Chesterfield Co. line, on S.C. 9, S.C.,
     26-V-75.  D. A. Etnier, G. A. Schuster (1 male mmt.,  1 larva).
                                     53

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Tennessee:
     Walker Br. Trib. to Tellico R., on Tn. 210, Monroe Co., Tn., 20-VI-74. ">
     D. A. Etnier, G. A. Schuster (1 male mmt.); Little Pigeon R., Maple Br.
     Rd., off Tn. 2421, near Richardson Cove, Sevier Co., Tn., 10-VII-74.
     C. A. Schuster, G. A. Schuster (1 male mmt., 4 larvae); George Cox
     Property, Webb Mtn., Sevier Co., Tn., 17-IV to 17-V-75.  G. A. Schuster
     (many adults); Little R., at U.S. 411, Blount Co., Tn., 23-IV-75.  C. A.
     Schuster and G. A. Schuster (1 male mmt.); Conasauga R., at Tn. 74,    ;i
     Bradley Co., Tn., 4-V-75.  D. A. Etnier and G. A. Schuster (1 male mmt.);
     Hurricane Cr., 2 mi. N. of Tn. 62, Fentress Co., Tn., l-V-75.  D. A.
     Etnier, G. A. Schuster (2 male mmts.); Little R., 2 mi. above Townsend,
     Blount Co., Tn., on Tn. 73, 14-VII-75.  M. Hughes, ₯. Dickinson, G.
     Schuster (several larvae); Abrams Cr., Great Smoky Mtn. Nat. Pk., Blount
     Co., Tn., 26-VII—75.  C. A. and G. A. Schuster (1 male mmt., several
     larvae); Stream at Montgomery Bell St. Pk. Entrance on U.S. 70, Dickson;
     Co., Tn., 2-V-75.  D. A. Etnier, G. A. Schuster (1 male mmt.); Little
     Pigeon R., 1 mi. S. of Richardson Cove, on Co. Road 2421, Sevier Co.,
     Tn., 21-VI-75.  G. A. Schuster (many larvae); Little R., in Townsend,  ..
     Blount Co., Tn., 19-VI-75.  M. Hughes, W. Dickinson, G. A. Schuster
     (15 larvae); Obed R., at Genesis Rd., Cumberland Co., Tn., 26-V-75.
     D. A. Etnier (1 male and 1 female mmt., 1 larva); Caney Fork R., 1.3
     mi. E. of Pleasant Hill, Cumberland Co., Tn., 14-IV-76.  G. A. Schuster
     (1 male mmt.) .

Wisconsin:
     Lowes Cr., 1 mi. W. of State St., Eau Claire, Eau Claire Co., Wi.,
     28-VIII-75.  D. A. Etnier (3 male mmts.).

Distribution

     Symplvitopsyche sparna is widely distributed in eastern North America.
Published records include the following states and provinces:  Georgia,
Kentucky, Maine, New York, Michigan, Minnesota, North Carolina, Nova Scotia,
Ontario, South Carolina, Tennessee, Virginia, West Virginia, and Wisconsin.
Unpublished records included here are from Alabama.

Biology

     This species has been collected widely over the southeastern United
States.  In this part of the country, S. spama seems to be ubiquitous.  It
has been taken in a variety of habitats ranging from first order streams to
large river situations.  In this respect S. spama seems to have the widest
range of tolerance to environmental conditions of[any Symphitopsyahe species
collected by the authors;  It is commonly collected;,in small, sluggish,
organically rich streams as well as fast, clean, cold trout-stream-type
waters.                                           ;::

     Its northern range, however, is more puzzling.  It has a rather spotty
distribution often commonly encountered and just as often absent altogether.
A good example of this situation occurs in Illinois, a state which has been
extensively collected (Ross, 1944) and one which is almost devoid of this
species (JJnzicker, personal communication).

                                     54

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     Collection records  indicate  this  species emerges  from April to September
 and  is probably univoltine.
                        Symphitopsyche Ventura  (Ross)
                                   (Fig. 27)

     Symphitopsyehe Ventura  (Ross), 1941.  Trans. Am. Ent. Soc., 67:92  (Type
locality:  Costellow Lake, Algonquin Pk., Ont. Fish. Res. Board, 10-VI-39.
W. M. Sprules).

     As Hydropsyche Ventura:  Ross, 1944.  111. Nat. Hist. Suv. Bui., 23:294;
Blickle and Morse, 1966.  Me. Ag. Exp. Stat. Tech. Bui., 24:6; Etnier, 1973.
J. Ga. Ent. Soc., 8:273.

Description

     Head capsule length, 1.20 to 1.28 mm; width, 0.99 to 1.10 mm.  Posterior
one-fifth of dorsal surface of head yellow; rest of head dark brown.  Lateral
aspects of head yellow; venter of head yellow with brown pigmentation on
stridulatory surface and geneal suture.  Genae covered with short, brown
setae dorsolaterally.  Frontoclypeus with such setae scattered over sclerite.
Labrum dark brown covered with short, black setae; laterally margined with
long, yellow setae.  Mandibles dark brown and edged in black.  Thoracic
sclerites golden to dark brown; covered with thin, short, brown hairs.  Each
notum also covered with short, golden spine-like setae which are not very
conspicuous.  Abdomen straw colored and covered dorsally and ventrally with
short, black hairs.  Each abdominal segment with numerous club hairs which
are also found below the pleural gills.  Dorsal sclerite of anal legs covered
with short, black hairs.  Venter of anal legs also covered with such hairs.

Diagnosis

     This species may be separated on the basis of the frontoclypeus color
pattern.  The frontoclypeus is unicolored dark brown rather than black.   It
may be confused with the inconspicuously spotted S.  sparna.  However, a
number of characters may be utilized to separate these two species.  First,
the abdominal gills of S,  Ventura are not as profusely branched as in S.
sparna.  Second, S. sparna has many conspicuous, thick short, black setae
on the thoracic nota; on the nota of S.  Ventura similar- setae are not as
numerous nor are they as thick and conspicuous.   Third,  the club hairs on
the abdomen of S.  Ventura are much more numerous and conspicuously thicker
than those of S. sparna. -Also, numerous club hairs are found below the
pleural gills in S. venturtt but not in S.  sparna.  Fourth, the head of S.
sparna, except for the typical three light spots on the frontoclypeus, is
unicolored; whereas, the dorsal surface of the head of S.  Ventura is much
darker than the lateral surfaces the the frontoclypeus is unicolored.

Material Examined

Newfoundland:
     Big Falls, Newfoundland, 28-VI-66.   D.  R.  Smith (adult males;  USNM)
collection).

                                      55

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New York:
     Bear Brook near Blue Mtn. Lake, Adirondack St. Pk.,  N.Y., 19-VT-41.
     Frison and Ross (adults; INHS collection); Cedar R.  near Indian Lake,
     Adirondack St. Pk., N.Y., 20-VI-41.  Frison and Ross (adults; INHS
     collection).

Tennessee:
     English Cr., at Carson's Springs near Newport, Cocke Co., Tn., 3-8-VI-
     46.  Mike Wright  (adults; INHS collection); Lost Cr., on waterfalls on
     Co. Rd. 4448, 6 mi. S.E. of Sparta, White Co., Tn.,  14-IV-76.  G. A.
     Schuster (2 males and 6 female mmts.); Anderson Cr., trib. to New R.,
     1.3 mi. S.S.W. of Montgomery Jet., W. of Co. Rd. 2344, Scott Co., Tn.,
     l-VII-76.  T. Talak (1 male mmt.); Jake Branch, at Tn. 63, on Scott/
     Campbell Co. line, 2-VII-76.  T. Talak (1 male mmt., 2 larvae).

Virginia:
     Gilletts Run, 10 mi. N. of Warm Springs, Bath Co., Va., 17-VII-71.
     A. B. Gurney (adult males; USNM collection).

Distribution

     Little is known of the distribution of this rarely collected species.
scattered collections have been reported from the following regions:
Newfoundland, New York, Ontario, Tennessee, and Virginia.

Biology

     The largest collection of metamorphotypes is from Lost Creek in White
County, Tennessee.  At this locality Lost Creek is a 60 degree to 90 degree
waterfall with moderate to fast currents.  The depth averages less than 6
inches and the width is approximately 30 feet.  The shore vegetation is
mixed deciduous which  produces cover.  The bottom consists primarily of
bedrock with some loose rocks of medium size near the upper portions of
the falls.  The substrate is lustily covered with moss.  No larvae were
collected here; however, the pupae were found attached to the moss growing
in the fastest water.  Since most of the substrate was bedrock, collections
had to be made by feeling for pupal cases or larval retreats.  This may be
the reason no larvae were collected since it is very difficult to obtain
access into the small  cracks and crevices of the bedrock  in which the
larval retreats are often located.

     Other larval caddisflies collected at Lost Creek  included:  Micrasema
sp., Goevita t>etten-i,  Diplectvona modesta, Oropsyohe ;sp., Hydropsyche  sp.
 (depvavata group) .                                 - - •'-

     According to available collection records, S. Ventura emerges  from
early April through September.
                                      56

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                        Symph-Ctopsyahe piatY"Lx (Ross)
                          (Presumed Larva, Fig. 28)

     SympTii-bopsyche piatrix (Ross), 1938.   111.  Nat. Hist. Surv. Bui. (Type
locality:  Greer, Missouri, 28-111-37, at spring near town.  T. H. Prison).

     As Hydropsyche p-iatvix:  Ross, 1944.  111. Nat. Hist. Surv. Bui., 23:88,
96, 97, 294; Corbet et al., 1966.  Can. Etn., 98:1290; Unzicker et al., 1970.
J. Ga. Ent. Soc., 5:171.

     These larvae have not been definitely associated with the adult form.
On the basis of having been collected near the type locality of S. p-Latrix
and because all other SympTi-itopsyche larvae collected in the area can be
recognized, it is presumed that these larvae are actually S.  piatrix.

Description

     Head capsule length, 1.25 to 1.30 mm; width, 1.05 to 1.10 mm. Head
predominantly golden with some dark brown pigmentation.  Frontoclypeus with
four rectangular brown bars arranged in center of sclerite.  One bar ante-
rior, and one posterior to two lateral bars.  This arrangement produces a
yellow spot located centrally.  Head capsule also with dark brown transverse
stripe across the width of the head at posterior insertion of frontoclypeus.
Genae covered dorsally and laterally with small,  clear setae; located ante-
rior to transverse brown stripe.   No pigmentation around eye or on venter
of head.  Labrum golden, covered with similarly colored setae dorsally;
laterally with long fringe of yellow setae.  Nota light brown to straw
colored; each covered with very small, clear setae.  All leg segments light
brown to straw colored and liberally covered with small, clear setae.
Prosternum golden with black markings posteriorly; poststernal plates solid
and golden in color.  Abdomen golden brown, covered with short, black setae.
Club hairs, if present, inconspicuous.  Dorsal sclerite of anal legs covered
with very thin, golden setae.  Ventral surface of anal legs with large
heavily sclerotized spine-like setae, especially concentrated anteriorly.
These setae at least as heavy as those on ventral sclerites of abdominal
segment IX.

Diagnosis

     The larvae available for study for all characteristically light colored,
much like S. sparna and S. macleod-i.  However, it is easily separated from
these two species on the,;basis of head color pattern  (Fig. 28) as described
above.  Furthermore, this, larva can be separated from all other Symphito-
psyche species by the following characters:  (1)  Few or no club hairs are
present on the dorsum of the abdomen; if they are present, they are incon-
spicuous.   (2) Heavy spines are located on the ventral surface of the anal
legs.  These spines are at least as large and as heavily sclerotized as
those on the ventral sclerites of the ninth segment.  No other Symphitopsyche
species possesses such large and conspicuous spines on the anal legs.
                                      57

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Material Examined

Arkansas:
     Down stream from Mammoth Spring, 40 yards N. of jet. with Spring R.,
     in Mammoth Spring, Fulton Co., Ar., 25-IX-76.  H. ₯. Robison (14
     larvae).

Missouri:
     Head waters of Current R., 15 mi. S.E. of Licking in Montauk St. Park,
     Dent Co., Mo., l-IV-67.  G. L. Harp (3 larvae); Seven Springs, head
     water of Current R., Montauk St. Park, Dent Co., Mo., 2-IV-67.  G. L.
     Harp (9 larvae).

Distribution

     Symphitopsyche piatvix is apparently restricted to spring-type habitats
in the Ozark Mountains of southern Missouri and Arkansas.  Only one record,
that of Corbet et al. (1966), lists the species from other than this
vicinity.  They record it from the St. Lawrence River in Quebec, but we are
inclined to believe it is a misidentification of S. wafkevi, a closely
related species.

Biology

     Other than to say that S. piatz"i-x is restricted to spring-like environ-
ments, we cannot add more concerning its biology.  It was described in 1938
by Dr. Ross and has been reported on only three other occasions, one of
which is probably in error.  The rarity of this species in collections and
reference to it in the literature cannot be considered an indication of the
abundance of this species.  It is more likely that this rarity is a product
of limited sampling by Trichopterologists in its geographical range.
Ebpefully, in the future, this species will be studied in greater detail.

                 Symphitopsyohe etn-Ler-L (Schuster and Talak)
                                  (Fig. 29)

     Symphitopsyche etnieri (Schuster and Talak), 1977.  J. Kan. Ent. Soc.
(Type locality:  Buffalo Springs Cr., Buffalo Springs, Grainger Co., Tn.).

Description

     Head capsule length, 1.40 to 1.50 mm; width 1.25 to 1.35 mm.  Head
dark brown excepting yellow area around eyes and large yellow spot on the
posterior lateral aspects of the genae.  Posterior:dorsal portion of genae
slightly lighter than rest of head with numerous dark brown circular mus-
cle scars.  Anterior one-third of frontoclypeus darker (almost black) than
rest of sclerite.  Short, brown setae present on dorsolateral aspects of
genae.  Scattered setae located at anterolateral corners and posterior angle
of frontoclypeus.  Labrum very dark brown, covered with black setae; late-
rally margined with long, yellow setae.  Mandibles dark brown, edged in
black.  Thoracic nota dark brown;  pronotum slightly darker than other two
                                     58

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nota.  Each nota covered with thin, short, black setae, and laterally edged
in black.  Mesonotum with wide, U-shaped mark posteriorly; metanotum with
short, black slash mark posteriorly.  Prosternum brown with anterior and
posterior .margins black.  Poststernal plates brown.  Anterior legs brown;
mes and metathoracic legs golden.  Each leg segment amply covered with
short, brown spine-like setae not unlike those of other species.  Abdomen
tan covered with moderate number of thin, short, black hairs.  Club hairs
on dorsum and below pleural gills only slightly thicker than other hairs.
Dorsal sclerite of anal legs covered with short black hairs as are the
ventral surfaces.

Variation

     No variation from the above description was found in the larvae studied.

Diagnosis

     This species may be separated from most other Symph-itopsyohe species on
the basis of head capsule coloration.  Only one other species, S. aThedra,
has similar pigmentation.  These two species may be reliably separated by:
(1) the presence of numerous circular dark brown muscle scars (Fig. 29) on
the dorsal posterior mesial portion of the head in S. etnlevl which are
absent in S. aUhedpa.', (2) the abdomen of 5. e~bnieT-i has considerably fewer
hairs especially on the ventral surfaces, and the hairs are thinner and not
as long as those of S. alhedra; (3) the club hairs are more slender and less
conspicuous in S. etn-ieT-i-; (4) in general, S. e-bn-Lern, is less robust and
smaller than S. alhedra.  The pigmentation of S. etnieT-i is not as dark as
that of S. alhedra\ also, the sclerites, especially those of the head, are
not as shiny.

Material Examined

Tennessee:
     Buffalo Springs Cr., 100 yds. W. of jet. Co. Rds. 2479 and 2480, Buffalo
     Springs, Grainger Co., Tn., 13-IX-76.  T. Talak (23 male and 17 female
     mmts., 60 larvae); Buffalo Springs Cr., 100 yds. W. of jet. Co. Rds.
     2479 and 2480, Buffalo Springs, Grainger Co., Tn., 18-IX-76.  T. Talak
     (5 adult males, 20 adult females).

Distribution

     The only distribution records for this species are those given above.
These are from the typeijlocality, Buffalo Springs Creek, in the community
of Buffalo Springs, Grainger County, Tennessee.  The stream is 100 meters
west of the junction of county roads 2479 and 2480.  This is approximately
3.3 miles southeast of the junction of county road 2480 and U.S. 11W.
Buffalo Springs is approximately 4.0 air miles southwest of Rutledge,
Tennessee, which is the county seat.
                                     59

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Biology

     Knowledge regarding the biology and emergence times of this species is
scant since it has been collected on only two occasions.   However, the
following can be stated regarding the type locality which may be indicative
of the preferred habitat of this species.

     Buffalo Springs Creek begins as a spring approximately 1,000 meters
upstream from the type locality.  Below this, much of the spring is diverted
into a trout hatchery from which the effluent is returned to the stream.
The effluent has a high concentration of organic wastes.  Due to the origin
of the stream and the presence of the trout hatchery, the water temperature
is fairly constant throughout the year.  Symphitopsyohe etni-eri. was the only
SympTiitopsyche species present, and Chewnatopsyehe eampyla Ross was the only
other caddisfly collected at this site.

     Existing in a cold-water, spring-type habitat is not unique to this
species of Symphitopsyche.  Many species of the genus seem to exhibit an
adaption to cold water situations.  For example, species such as S. p-iatvix
Ross and S. maeleod-i Flint have been collected only in these types of
environments.

                                     60

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                            The Genus Hydropsyche

                           Depravata Species Group

                          Hydropsyohe betteni, Ross
                               (Fig. 1 and 30)

     Hydropsyohe betten-i Ross, 1938.  111. Nat. Hist. Surv. Bui. 21:146-147
 (Type locality:  Richard, 111., 28-V-36.  Ross).

     Hydropsyohe betteni:  Betten, 1934.  Caddis Flies N.Y. St., p. 188 (as
 incommoda, not Hagen); Ross, 1941a.  Tr. Amer. Ent. Soc., 67:85; Denning,
 1943.  Ent. Amer., 23:103-105; Ross, 1944.  111. Nat. Hist. Suv. Bui., 23:86,
 91, 93, 99-100, 294; Leonard and Leonard, 1949.  Occ. Pap. Mus. Zoo., Mich.,
 522:9; Etnier, 1965.  Ent. News, 76:146; Blickle and Morse, .1966.  Me. Ag.
 Exp. Sta. Tech. Bui., 24:6; Edwards, 1966.  J. Tenn. Acad. Sci., 49:120;
 Unzicker et al., 1970.  J. Ga. Ent. Soc., 5:171; Longridge and Hilsenhoff,
 1973.  Wise. Acad. Sci., Arts and Letters, 61:176; Resh, 1975.  Trans. Ky.
 Acad. Sci., 36:12.

 Description

     Head capsule length, 1.40 to 1.50 mm; head capsule width, 1.25 to 1.40
 mm.  Entire head very dark brown to black except for yellow area around eye;
 occasionally with pair of inconspicuous pale spots on anterior portion of
 frontoclypeus.  Numerous bristle-like, black setae on top and sides of genae;
 similar setae on anterolateral corners and across front margin of frontocly-
 peus.  Area at posterior angle of frontoclypeus slopes downward.  Labrum dark
 brown to black with numerous black setae; laterally fringed with long, yellow
 setal brush.  Mandibles dark brown to black.  Nota dark brown, laterally
margined in black, each notum with many bristle-like setae.  All legs dark
brown, abundantly adorned with black setae and gold spine-like setae.
Prosternum bronze, anteriorly and posteriorly edged in black.   Prosternal and
mesosternal plates dark brown fairly well sclerotized.  Abdomen brown; covered
with numerous scale hairs; equally abundant on all abdominal segments.
Minute spines present on dorsum of all abdominal segments.  Anal legs lacking
large, heavily sclerotized spine-like setae, but with minute spines present
on venter of last two abdominal segments.

Variation

     Hydropsyehe betten-i larvae are typically black-headed with no light
markings except those around the eyes.  However, some populations do have
 individuals which may exhibit a pair of medium-sized spots on the frontocly-
peus.  These spots are only slightly lighter than background color and are
                                      61

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rather inconspicuous.  This condition is rather atypical, but one population
found consists primarily of such individuals.  This population occurs below
Lake Toxaway Dam on U.S. 64 in Transylvania County, North Carolina.

Diagnosis

     The larvae of this species most closely resemble those of H. depvavaba
and H. potomaoens'is, to which it is closely related.  At this time no reliable
characters have been discovered to separate these three species.  Hydropsyche
betteni- may at times, however, be separated from the other two species on the
basis of its range, since the ranges of H. depx>avata and H. potomaeensis are
much more restricted.

Material Examined                  ,

Kentucky:
     Silver Cr., on Ky. 876, 5..0 mi. W. of Richmond, Madison Co., Ky., 7-VII-
     76.  C. A. and G. A. Schuster (2 male mmts., several larvae); Silver Cr.,
     on Ky. 876, 5.0 mi. W. of Richmond, Madison Co., Ky., 13-VI-76.  G. A.
     Schuster (several larvae).

North Carolina:
     Panther Cr., at Fines Cr. exit on 1-40, Haywood Co., N.C., 24-V-75.  G.
     A. Schuster, D. A. Etnier (2 male mmts.); Shoal Cr., on U.S. 441, 1.0 mi.
     S. of Cherokee, Jackson Co., N.C., l-V-76.  G. A. Schuster (2 male mmts.,
     4 larvae); Cullasajai R., 6.0 mi. W. of Highlands, Macon Co., N.C., 19-VI-
     76.  G. A. Schuster (several larvae); Cullasaja R., just below dam at
     Highlands, Macon Co., N.C., 27-VI-76.  D. A. Etnier (several pupae and
     larvae); Below Toxaway Dam, 10 mi. E. of Cashiers, Transylvania Co., N.C.,
     on U.S. 64, 2-VIII-76.  C. A. and G. A. Schuster (many mmts. and larvae).

Tennessee:               ',
     Unnamed Cr., 1.0 mi. from Johnson Bible College, Knox Co., Tn., 12-IV-75.
     D. and L. Etnier, G. A. Schuster (2 male mmts.); Turkey Cr., Loudon Co.,
     Tn., ll-IV-75.  D. A. Etnier (1 male mmt.); Flat Cr,, Morgan Co., Tn.,
     26-IV-75.  D. A. Etnier (several mmts. and larvae); Gap Cr., off Co. Rd.
     2586, 2.0 mi. S. of Johnson Bible College, Knox-Sevier Co. line, 16-IV-
     75.  G. A. Schuster (1 male mmt.); .Hurricane Cr., 2.0 mi. N. of Tn. 62,
     Fentress Co., Tn., l-V-75.  D. A. Etnier, G. A. Schuster (3 male mmts.);
     Caney Fork R., on unnamed Co. Rd., 2.0 mi. N.E. of Pleasant Hill,
     Cumberland Co., Tn., 23-VII-75.  G. A. Schuster (1 male mmt.); Little
     Emory R., near Wartburg, Morgan Co., Tn., 26-IV--75.  D. A. Etnier (1
     adult male, 1 male mmt., several larvae); Clean .Cr., at Fentress and
     Cumberland Co. line:on .Tn. 62, l-rV-75. . D. A.: Etnier, G. A. Schuster
     (1 male mmt., several larvae); Bull Run Cr., on Tn. 162 just S. of Oak
     Ridge, Knox Co., Tn., 8-IV-76.  D. A. Etnier, :G. A. Schuster (1 male
     mmt., 3 larvae); Unnamed Cr., at University of Tennessee Arboretum, Oak
     Ridge, Morgan Co., Tn., 6-IV-76.  D. A. Etnier (several male mmts. and
     larvae).

Virginia:                         ,  •
     Small Trib. to James R., 0.4 mi. E. of Va. 615 on Va. 621, Powhattan Co.,
                                      62

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      Va., 13-VI-75.  D. A. Etnier, W. C. Starnes (several mmts. and larvae).

 Distribution

      This species has been recorded from the following states and provinces:
 Arkansas, Georgia, Illinois, Indiana, Kentucky, Maine, Michigan, Minnesota,
 New Hampshire, Ohio, Ontario, Pennsylvania, Tennessee and Wisconsin.  With
 the additional new records from North Carolina and Virginia as listed above.

 Biology

      As stated above, this species is very common in small, warm-water streams.
 It is often found in very large numbers, much like H.  orris in large rivers.
 H. bet-bent seems to be one of the most resistant Hydropsyahe species to or-
 ganic pollution.  Its ability to cope with and use the high concentrations of
 organic materials may contribute to the high numbers of individuals in the
-populations collected.  It has been found that, where H.  betteni occurs, it
,is usually the most predominant caddisfly present.   It is often the only
 Hydropsyohe species in the stream, but it is not uncommon for other species
 to occur with H. betteni.   The species most often collected with H.  betteni
 is Symphitopsyohe sparna which also seems to be resistant to high concentra-
 tions of suspended materials.

      The emergence period of H.  betteni is from April  to  September.

                          Hydropsyohe depravata Hagen

      Hydropsyche depravata Hagen,  1861.   Syn.  Neur.  N.  Am., p.  290 (Type
 locality:   "Dalton,  Ga.  (Osten Sacken)").

      Hydropsyohe depravata:   Hagen, 1864.   Verh.  Zoll.  Bot. Ges.,  14:822;
 Banks,  1892.   Tr.  Amer.  Ent.  Soc., 19:367;  Ulmer, 1905.   Z. Insbiol.,  1:68;
 Ulmer,  1907.   Gen.  Ins.,  60:71;  Banks,  1907.   Cat. Neur.  Ins. U.S.,  p.  47;
 Betten,  1934.   Caddis Flies N.Y.  St.,  p.  187-188; Milne,  1936.   Stud.  N. Am.
 Trich.,  3:70,  72,  73;  Banks,  1936.  Psyche,  43:129;  Ross,  1938c.   Psyche,
 45:17;  Brimley,  1938.   Ins.  N.  Carolina,  p.  251;  Ross,  1944.  111. Nat.  Hist.
 Surv. Bui.,  23:15,  91,  93,  100,  294;  Edwards,  1966.  J. Tenn. Acad.  Sci.,
 41:120;  Resh,  1975.   Trans.  Ky.  Acad.  Sci.,  36:12.

 Variation

      The larvae  of  this  species  are generally  dark-headed  like H.  betteni
 with  light  areas only around the eyes.  However,  one population  has  been
 collected from the  Stones: River  near Nashville,, Tennessee,  which is  much
 lighter  in  color.   The head  capsule of  these specimens  is  rich brown rather
 than  black with  lighter  areas behind the  eyes.

 Diagnosis

      Hydropsydhe depravata larvae  and metamorphotypes have been  collected
 from  a number  of different localities; however, to date this species cannot
 be separated from larvae of.-H. betteni and H. potomaaensis  (see Fig. 30).  No
                                      63

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reliable characters have been found to consistently differentiate the above  ,1;
three species from one another.

Material Examined

Tennessee:
     Bennett Cr., on Bennett Cr. Rd., at Bennett Cr. Baptist Church, Rnox Co.,
     Tn., 16-IV-75.  G. A. Schuster (1 male mmts., numerous larvae); Stock Cr(.,
     off Tipton Station Rd., off U.S. 441, Knox Co., Tn. , 16-IV-75.  G. A.   :::
     Schuster (2 male mmts., several larvae); Turkey Cr. ,, off Northshore Dr.,
     Knoxville, Knox Co., Tn,, 17-IV-75.  D. A. Etnier  (1 male mmt.); Small
     Cr., 1.0 m±. S.. of Johnson Bible College, Knoxville, Knox Co., Tn., -12- "
     IV-75.  D. A. and L. Etnier, G. A. Schuster (1 male mmt., several larvae),
     Caney Fork R., 2.0 mi. N.E. of Pleasant Hill on unnamed Co. Rd., Cumber^
     land Co., Tn., 14-IV-76.  G. A. Schuster (many larvae, several mmts.).

Georgia:
     Below dam on spring at jet. Ga. 2 and Ga. 223, Murray Co., Ga., 25-IV-76.
     G. A. Schuster, D. A. Etnier, J. Williams (3 male mmts., many  larave).

Distribution

     Hydvopsyche depravata is known from the following  states:  Indiana,
Kentucky, North Carolina, and Tennessee.  The above collection locale from
northern Georgia represents a new distribution record.  Its range is much
more restricted than that of H. betteni,.

Biology                  '.

     This species, like H. betteni, lives typically in  small, warm-water
streams which have high amounts of organic material.  The larvae are found in
the riffle areas of these streams on medium-sized rocks.  As with H. betteni,
when H, depravata is present, it occurs in large numbers.  On occasion, H.
depvauata and H. betteni, metamorphotypes have been  collected in the same
stream at the same time; however, the two larvae are indistinguishable from
one another.

                       Hydvopsyahe potomaoensi-s Flint

     Hydvopsyche potomacens-is Flint, 1965.  Proc. Ent.  Soc. Wash.,  67:169-171
(Type locality:  Virginia, Highland Co., bridge on Rt.  220 over E.  Fk. Potomac
R., 18-20-V-63.  W. D. Field and 0. S. Flint, Jr.). r

Diagnosis                                          -

     Hydvopsydhe potomacensi^s has been associated, but no reliable  characters
have been found to separate it from the larvae of H. betteni and H. depravata
(Fig. 30).
                                      64

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Material Examined

Virginia:
     North R., at the bridge at jet. of Va.
     Crawford, Rockingham, Co., Va., 17-VIII-75.
     10 larvae).
11 and Va. 727, out of Mt.
      G. W. Wolfe (2 male mints.,
Distribution
     This species was previously known only from the type material which was
collected from the East Fork of the Potomac River.

Biology

     Because this species has been collected only twice, very little is known
of its biology.  The river in which the larvae were collected is a medium,
warm-water river similar to the type of habitat in which H. depravata and H.
betten-i- have been commonly collected.

     Based on the type material and the pupal collections, it is surmised that
the emergence period is at least May through August and, in all probability,
may be longer than this.

                          Hydropsydhe elissama Ross
                              (Fig. 31A and B)

     Hydropsyche elissoma Ross, 1947a.  Trans. Am. Ent. Soc., 73:137-138 (Type
locality:  Mossy Cr., Perry, Ga.), 31-111-45.  P. W. Fattig).

Description

     Head capsule length, 1.10 to 1.20 mm; head capsule width, 1.05 to 1.15
mm.  Head on top mostly dark brown; frontoclypeus with a pair of large, dia-
gonal, tear-shaped yellow marks.  Area around epicranial stem yellow; brown
blotch laterally to this consisting of several darker oval muscle scars.
Similar smaller, dark blotches of muscle scars lateral to these.  Area around
eye and behind it yellow reaching dorsally posteriad to form rough, L-shaped,
light area.  Venter of head bronze, not as dark as top of head,  with few
scattered, slightly darker muscle scars.  Head from lateral view distinctly
deeper posteriorly than anteriorly.  In dorsal view head wider anteriorly,
sides of head with slight constriction centrally.  Frontoclypeus at posterior
angle distinctly thickened into large, low and rounded tubercle, best observed
in lateral view.  Top and sides of genae with numerous bristle-like, black
setae; present on the frontoclypeus only on the anterolateral corners.   Labrum
brown; adorned with black and yellow setae; laterally fringed with long,
yellow, setae.  Mandibles golden with black margins; pronotum slightly darker
than other two nota.  Pronotum and mesonotum with numerous stout, bristle-
like setae each in a brown pocket giving nota freckled appearance; metanotum
with very few of these.  Prosternum clear yellow with black posterior margin;
prosternal and mesosternal plates clear yellow, not heavily sclerotized.
Legs yellow, anterior pair slightly darker than others.  All legs adorned with
numerous spine-like and bristle-like setae.  Abdomen brown with numerous scale

                                      65

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hairs posteriorly decreasing in number and size anteriorly.  Minute abdominal**'
spines on dorsum of all segments decreasing in size and number posteriorly.  '>'
Venter of anal legs without large, sclerotized spine-like setae, but with    :--
minute spines that are on venter of both segments VIII - IX as well.         ;SJi
                                                                            : ;ci'
Diagnosis                                                                   ~'~'',

     This species can be most easily distinguished from others on the basis  !»
of the large, rounded tubercle (Fig. 31B) on the posterior angle of the fronto-
clypeus.  The color pattern is very similar to that of H. deealda which also.•: :
has two large tear-shaped marks on the frontoclypeus.  But in the case of alle
other species, H. deealda lacks the posterior frontoclypeal tubercle.       ::

Material Examined                                                            ;

Georgia:                                                                     :
     Beaver Cr., 1.0 mi. S. of Roberta, Crawford Co., Ga., 5-IX-76.  G. A.
     Schuster  (1 male mmt., 10 larvae).

Distribution

     This species was previously known only from the type material .described.
by Ross  (1947a).  The larval material examined was collected approximately  20
miles from this locality.

Biology                  ;

     Hydropsyche eli,ssoma  is one of the least known Hydropsyche  species.   The
stream from which the above larvae were collected was approximately 1.2 meters
wide and 0.6 meters  deep and approached a black-water condition.  The bottom
was dominantly sand  with some medium-sized rocks in a short riffle area.   No
larvae were  collected in the riffle on rocks; all were  collected in submerged
branches and logs into which they bored larval  retreats.   The  larvae themselves
were in  canals in the wood, and around each hole was a  rather  short net.   The
larvae were  collected by dismantling  the wood;  the pupa was found in the wood
itself.  The pupal  case was distinctly pliable  and consisted of  mostly of
secreted material with  some wood  fibers  incorporated.   Collected with H.
elissoma were  several H. deealda  larvae.

     Little  can be  said  of the emergence  period of H. elissoma since it has
been collected so infrequently.   The  type material was  collected in March  which
suggests  that  this  species emerges  at least  from March  until September.

                          Hydropsyche deealda Ross
                                  (Fig. 32)       ..„.;.;

     Hydropsyehe deealada Ross, 1947a. .Trans. Am.  Ent.  Soc., 73:138-139 (Type
locality:  Beaver Cr., 5 mi. S.E. of Roberta, Ga., 6-IX-45.  P. W. Fattig).

     Description

     Head capsule length, 1.30 to 1.45 mm; head capsule width,  1.25 to 1.40 mm.

                                      66

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 Head predominantly straw-colored with large areas of brown and numerous small,
 oval, dark brown to black muscle scars.  The frontoclypeus brown with pair of
 large, tear-shaped, yellow spots anterolaterally.  Frontoclypeus with numerous
 small dark brown to black pits.   Genae mostly straw-colored with numerous dark
 brown, oval muscle scars dorsally and laterally;  venter of genae slightly
 darker and also with muscle scars especially posteriorly.   Genae with short,
 bristle-like,  black setae on top and sides,  interspersed with many clear, short,
 spine-like setae.   Posterior half of frontoclypeus with many thin, short, clear'
 setae.  Head from lateral view distinctly sloping; deeper  posteriorly.   In
 dorsal view, head wider anteriorly.   Sides of anterior half of head bulging
 outward.   Nota same color as head and laterally margined in black.  Pronotum
 and mesonotum with numerous bristle-like,  black setae and  short,  clear  spine-
 like setae in dark brown pits giving sclerites freckled appearance.   Each
 notum with several larger,  dark  brown,  round muscle scars  on lateral aspects
 of  sclerites;  pronotum with about twice as many as other nota.  Prosternum
 yellow with black posterior margin;  prosternal and mesosternal plates yellow
 and weakly sclerotized.   Legs yellow;  anterior pair slightly darker;  all  legs
 covered with many  spine-like setae and  long,  black setae.   Abdomen brown;
 covered with numerous  scale hairs posteriorly;  decreasing  in sized and  number
 anteriorly.  Minute spines  on dorsum of all  abdominal segments  VIII  - IX.
 Anal legs  lacking  large,  heavily sclerotized  spine-like setae on  ventral
 surfaces.

 Diagnosis

      This  species  is closely related  to H. eHssoma and resembles  most  closely
 that  species.  These two  species  differ from  other species  in the  depravata
 group  In that  they  lack a coal-black head  and nota.  H. decalda can be  recog-
 nized  by the color  pattern  of  the head, 'especially the  two,  large, tear-shaped,
 diagonal marks on the  frontoclypeus, the small, brown pits  on the  frontoclypeus,
 and  the numerous brown, oval muscle scars  on  the top, side,  and bottom  of the
 genae.  This color  pattern  is not too unlike  that  of .#. elissoma except that
H. eltssoma has a much darker  frontoclypeus and not as many muscle scars on
 the genae.  The single most distinguishing character between  these two  species
 is the lack of the posterior frontoclypeal tubercle in .H. deoalda.  It can be
readily observed in lateral aspect in E. -eltssoma.    •.   •

Material Examined
Georgia:
     Beaver Cr., 1.0 mi. S. of Roberta, Crawford Co., Ga., 15-IX-76.
     Schuster (3 larvae).uu
G. A.
Louisiana:                        . ,  ., ..^. ..-. .  • :;  .......
     Little Bayou Pierre, Natchitoches Par., La., 29-XII-73.  J.  A.  Louton (6
     larvae); Little Bayou Pierre, Natchitoches Par.,  La., 17-111-74.   J.  A.
     Louton (11 larvae);  Little Bayou Pierre,  Natchitoches Par.,  La.,  5-IV-74.
     J. A. Louton (3 larvae);  Little Bayou Pierre, Natchitoches Par.,  La., 22-
     V-77.  D. A. Etnier, U.  T. Regional Faunas Class  (several larvae, 3 male
     mmts.);
                                      67

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     Big Cow Cr. at Hwy. 63, Newton Co., Tx., 22-V-77.  D. A. Etnier, U.  T.
     Regional Faunas Class (several larvae, 1 male mmt.).

Distribution

     Previously this species was known only from the type locality from which
three H. decolda larvae were collected.  The above records from Little Bayou
Pierre, from which adults have been taken as well, and Texas constitute a
significant range extension of this species.  If more work were to be done in
the south, it may be found that this species is fairly widespread.  Its rarity
in collections may well be due to lack of strenuous light-trap collecting in
the south rather than it being naturally uncommon.

Biology

     Very little is known of the biology of this species.  The type locality,
Beaver Creek, as described under H. elisscma, is a rather small stream with a
predominantly sand bottom.  The larvae collected here were removed from sub-
merged logs and branches into which they bored out holes for larval retreats.
The Little Bayou Pierre locality, on the other hand, is slightly different
from most Louisiana streams in that the bottom consists of rocks and gravel
with large riffle areas.  The larvae at this locality were taken from the
rocks in the riffle areas.

     Little is known of the emergence of H. decalda.  Some adults were taken
at Little Bayou Pierre  in March, and the type material was collected in
September.

                          Hydropsyche oaTol-ina Banks
                              (Fig. 33A and B)

     Eydvopsyohe Carolina Banks, 1938.  Psyche. 45:77-78  (Type locality:
"North Carolina").

     Eydvopsyche carol-inai  Ross, 1944.  111. Nat. Hist. Surv. Bui., 23:294.
Flint, 1966.  Proc. U.S. Nat. Mus., 118:375.

Description

     Head capsule length, 1.10 to 1.25 mm; width, 1.00 to 1.15 mm.  Except for
small, circular, yellow area around eyes, entire head black.  Head with a broad,
concave frontoclypeus set off by an extensive arcuate carina.  Frontoclypeus
with greatest width just anterior to eyes.  From widest point to posterior
angle, frontoclypeal margins straight.  Anterior margin of frontoclypeus
straight.  Genae covered dorsally and laterally with short, stout, black
bristle-like setae; frontoclypeus devoid of such setae.  Labrum black, covered
with similarly colored setae, and margined laterally with brush of long, yellow
setae.  Mandibles black and typical.  Thoracic nota lighter in color than head;
dark brown dorsally, golden laterally.  Each nota edged in black laterally and
covered with short, stout, golden spine-like setae.   Leg  segments
                                      68

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 similar in coloration to  nota.   Each coxa and femur with proximal and distal
 borders trimmed in black.   Setation on legs not unlike that  of other Hydro-
 psyche  species.  Prosternum golden brown edged anteriorly and posteriorly in
 black.   Poststernal plates  rectangular,  dark brown;  mesosternal plates tri-
 angular; metasternum lacking plates.   Abdomen beige covered  with short,  thin,
 black setae.   Dorsum of each segment  with scale hairs  with greatest  number on
 last  several  segments; segments  I  - II with the least.   Dorsal sclerite of
 anal  legs  covered with black setae; ventrally anal  legs covered with numerous,
 short,  black  setae.

 Diagnosis

      This  small and distinctive  species  is  easily recognized on the  basis  of
 the presence  of a wide carina on the  head.   It is the  only known Hydpopsyohe
 larva which has such a carina and, with the present  keys available,  would  key
 to Uaovonema.   However, it  is unlikely that anyone  familiar  with Macponema
 would confuse the two genera.  There  are a  number of striking differences
 between H.  ccacolina and any of the Macronema species.   First,  the carina is
 much  more  offset in Macronema.   Second,  the carina  in #.  carol-ina is produced
 by the  suture joining the genae  and frontoclypeus, whereas, the carina of
 Maoronema  is  lateral to the frontoclypeus-genae suture.   Third, Macronema  spp.
 are typically pigmented a bright green with H.  Carolina having dark  brown  to
 black coloration.   Last, Macronema spp.  are at least two to  three times  larger
 than  the last  instar of H.  Carolina.

 Material Examined

 Georgia:
      Lakemont,  Ga.,  30-VI-39.  P. W.  Fattig (1 male, 3  females; INHS
      collection).

 North Carolina:
      Cullasaja  R.,  11.9 mi.  above jet. N.C.  28  and Bypass U.S.  441 and 23,
     Macon  Co.,  N.C., 9-V-76.  D. A.  Etnier (many larvae); Cullasaja R., 6 mi.
     W. of  Highlands, Macon  Co., N.C., 19-VI-76.  G. A.  Schuster  (4 larvae, 1
     male mmt.,  3  female mmts.); Cullasaja  R.,  6 mi. W.  of Highlands^ Macon
      Co., N.C.,  19-VI-76.   G. A. Schuster (41 males, 34  females); Oconaluftee
     R., at Cherokee, Swain  Co., N.C., 27-VI-76.  D. A.  Etnier  (1 larva).

Distribution

     Hydropsyche Carolina has been so rarely collected that little can be said
regarding its distribution.  The few collection records  available indicate
that it is  restricted to areas in and about the mountains of North Carolina
and Georgia.

Biology

     This species  is one of  the most poorly known and least understood of all
Hydropsyche species.  Our largest collections came from the Cullasaja River
near Highlands, North Carolina.   This river is impounded in Highlands with the
                                      69

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reservoir overflow making up most of the water volume downstream.  Due to
this, the water temperature is atypically warm for much of the year even though
the elevation is rather high (approximately 1,200 meters at Highlands).  Most
of the larvae as well as the largest known collection of H. Carolina adults
were taken at an elevation of about 1,000 meters.  The river here has moderate-
to-fast current with a depth of 0.5 to 1.5 meters and a width of 20 meters.
The shore vegetation is primarily Rhododendron with little aquatic vegetation
being present.  The bottom consists primarily of large, smooth boulders to
which H. Carolina larval retreats and pupal cases were attached.  The larval
retreat is not unlike that of other Hydropsyche species, but the pupal cases
were strikingly flimsy.  They were first thought to be Dolophilodes pupal
cases until they were later examined in the laboratory.  Collected sympatri-
cally with H. Carolina were the following Hydropsychids:  Diplectrona modesta
Banks; Cheumatopsyche campyla Ross, Hydropsyche betteni Ross, and Symphito-
psyche sparna (Ross).

     The few collections of H. Carolina indicate the species emerges in May
to June.

                             Cuanis Species Group

                           Hydropsyche cuanis Ross
                                   (Fig. 34)

     Hydropsyche cuanis Ross, 1938a.  111. Nat. Hist.  Surv. Bui., 21:147-148
(Type locality:  Wilmington, 111., 17-V-37, on Kankakee River.  Ross and Burks).

     Hydropsyche cuanis:  Ross, 1944.  111. Nat. Hist. Surv. Bui., 23:6, 87,
91, 93, 100-109, 294; Leonard and Leonard, 1949.  Occ. Pap. Mus. Zoo., Mich.,
522:9; Edwards, 1966.  J. Tenn. Acad. Sci., 41:120; Longridge and Hilsenhoff,
1973.  Wise. Acad. Sci., Arts and Letters, 522:176.

Description

     Head capsule length, 1.40 to 1.55 mm; head capsule width, 1.20 to 1.35 mm.
The following description was given by Ross (1944:100);  "Head and thoracic
sclerites bright brownish yellow, the head with an irregular, fine, reddish
brown pattern, the pronotum with fine, reddish brown speckling; legs yellow.
Frons almost flat, the apical margin straight.  Dorsum of  abdomen, especially
on the  seventh and eighth segments, with conspicuous flattened setae inter-
spersed among the simple appressed ones."  In addition to  the above:  Fronto-
clypeus covered with numerous, short, clear spine-like setae except for large,
shiny area anterior to pair of large, inconspicuous spots.  Dorsolateral
aspects of genae and pronotum with numerous such setae; mesonotum and metanotum
with smaller and fewer spine-like  setae.   Frontoclypeus, genae and nota lacking
long, thick, stiff, black setae.  Abdomen  with very few short, "simple appressed"
hairs dorsally; more below pleural gills than above;   Greatest concentration of
such hairs on venter of thorax.  Flattened hairs on dorsum more  like club  hairs
of Symphitopsyche than scale hairs, very sparse on segments VII  - VIII.  Minute
brown spines on dorsum of all abdominal segments.  Dorsal  sclerite of  anal
legs with long, brown setae; ventrally without spine-like  setae, with  short
setae.

                                      70

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  Diagnosis

       Hyfoopsyohe ouanis may be distinguished from other species in the genus
  by the following combination of features.  First, the sclerites of the larvae
;  are reddish brown with two large, but not very distinct, spots on the fronto-
  clypeus.   The genae also possess many smaller and irregular spots especially
  on the top and sides.   Second, the frontoclypeus, genae, and nota have
  numerous,  small, brown spine-like setae and lack heavy, stiff, black setae.
  Third,  the dorsum of the abdomen possesses numerous, minute spines,  with few
  flattened  setae and fewer short, thin appressed setae.   The flattened setae
  are most abundant on segments VII - VIII, but even here they are not numerous.
  The venter of the abdomen and thorax also possesses more short,  appressed
  hairs than does the dorsum of the abdomen.   Last, the anal legs  lack spine-
  like setae on the ventral surfaces.

  Material Examined

  Illinois:
       Kankakee R.,  at Wilmington, Grundy Co.,  II., 27-V-35.   Ross and Mohr (1
       male  mmt.;  INKS collection);  Kankakee R.,  at Wilmington,  Grundy Co.,  II.,
       10-IV-35.   Ross and Mohr (13  larvae;  INKS  collection).

  Distribution

       Ross  (1944)  indicates  that  the  range is  restricted to  Illinois,  Indiana,
  and Michigan,  but more  recent collection records  extend the range northwest
  to Wisconsin (Longridge and Hilsenhoff,  1973) and south to  Tennessee (Edwards,
  1966).

  Biology

       Ross  (1944:100) gave the following  notes on  the biology of H. ouan-Ls:
  "Most of our  Illinois records  of this species are from  various points along
  the Kankakee River;  in  addition  we have  taken it  from two other points in the
  extreme northeastern corner of the state.  The larvae are extremely abundant
  in swift rapids of the Kankakee  River at Wilmington, and here we have taken
  large flights of the adults.   In this locality the spring emergence during
 May is very heavy.  Adults continue to emerge later in  the year until August
  but never  in the large numbers that we have taken in May."

                            Soalca"is Species Group

                         -'  ; -Hydropsy ohe owls Ross
                         ;. •:'   (Fig. 35A and B)

      Hydropsyche oomuta Ross, 1938a.  111.  Nat.  Hist. Surv. Bui., 21:148,
 preoccupied by Martynov, 1909 (Type locality:   Hamilton, II., August  30,  1931.
 Ross and Mohr).

      Hydropsyohe orris:   Ross, 1938d.   Wash.  Ent.  Soc. Proc.,  40:121, new name;
 Ross,  1941.  Tr.  Amer.  Ent.  Soc., 67:86;  Denning,  1943.   Ent.  Amer.,  23:110,
 111,  118; Ross, 1944.  111.  Nat.  Hist.  Surv.  Bui., 23:5, 13,  86,  93,  95,  106-

                                       71

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107, 294; Leonard and Leonard, 1949.  Occ. Pap.  Mus.  Zoo., Mich.,  522:10;
Etnier, 1965.  Ent. News, 76:146; Unzicker et al., 1970.   J. Ga.  Ent.  Soc.,
5:171; Edwards, 1966.  J. Tenn. Acad. Sci., 41:120; Edwards, 1973.  Tex.
J. Sci., 24:504; Longridge and Hilsenhoff, 1973.  Wise.  Acad. Sci., Arts
and Letters, 61:176; Resh, 1975.  Trans. Ky. Acad. Sci.,  36:12.


Description

     Head capsule length, 1.35 to 1.50 mm; head capsule width, 1.30 to 1.40
mm.  Anterolateral margin of frontoclypeus with large conspicuous denticles.
Color pattern of head with clear, yellowish, V-shaped mark in middle of
frontoclypeus.  Lateral areas of this mark contiguous with clear area around
eye.  Anterior to V-shaped mark frontoclypeus with diamond-shaped black^area.
Coloration posterior to clear marking golden brown not as dark as anterior
section of- frontoclypeus.  Head without conspicuous black bristle-like setae,
but covered with small, slender, clear inconspicuous spine-like setae.  Head
in lateral view broadly rounded ventrally; dorsally flattened except for
raised teeth on frontoclypeus.  Laterally, large areas lacking dark coloration
posterior and ventral to eye.  Posterior  one-fourth of head capsule lacking
pigmentation.  Labrum straw colored; lateral hair fringes also straw colored.
Pronotum anterior edge almost lacking spine-like setae or hairs.  Pronotum
covered with very short spine-like  setae  that are same ground color, very
indistinct.  Pronotum laterally edged in  black  as is posterior margin.
Pronotum and mesonotum without hairs; metanotum with few hairs, very short
and slightly darker  than sclerite.   Thoracic sclerites and legs straw
colored.  Prosternum with black  spots on  anterolateral corners and black
bar on the posterior border.   Sclerites posterior  to prosternum S-shaped,
occasionally separated in center so that  each sclerite is divided.^ Sternal
plates on mesothorax wide and  slender with lateral margins projecting
anteriorad.  Ventral gills typically arranged.  Venter of anal legs covered
with  numerous  slender, golden  spine-like  setae.   Scale hairs very dense on
dorsum of abdominal segments  IV  - VIII.   No scale hairs below pleural gills.
Abdominal segment  III with scale hairs  spaced farther apart, not  as dense
and decreasing greatly on segment II, and present only on lateral aspects
of segment  I.   Minute spines  on  dorsum  of all abdominal segments;  greatest
concentration  on segments I - IV, becoming very small and inconspicuous
posterior to segment IV.


Variation                           ;

      Little variation of pattern exists either  geographically or  within
a given population.  Occasionally,  the  light, V-shaped mark may be broken
and produce a pattern similar to that of E.  bidens.,  but  the coloration of
 the lateral and ventral  aspect of the head rarely varies.
                                      72

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Diagnosis

     This species is most closely related to E. b-Cdens.  At both the adult and
larval level, these two species may at times be difficult to separate, as no
consistent morphological characters could be found to separate the larvae.
However, it was found that the color patterns of the head and ground coloration
of thoracic sclerites and legs can be used to reliably separate the two species.
Thoracic sclerites and legs of H. orris are straw colored to brownish yellow;
whereas, in E. bidens these bear a golden brown to dark brown coloration.
Differences in head capsule coloration are more pronounced.  Hydropsyche orris
bears a light, V-shaped mark (Fig. 35A) which is broken in the center in H.
bidens to produce two distinct, large spots (Fig. 36A) .  From dorsal view, the
entire posterior one-fourth of the head lacks pigmentation in H. owls.  In
contrast, there is dark pigmentation in E. ~bi,dens on either side of the suture
line connecting the right and left genae.  This gives the appearance of two
large, oblong, light spots at the posterolateral corners of the head.  In
lateral view of E.' orris, there are large areas where there is little pigmen-
tation (Fig. 35B) especially around and posterior to the eye.  Hydropsyohe
bidens, however, exhibits much dark pigmentation laterally (Fig. 36B) .  The
area around the eye is light, but posterior to this there is only a thin,
light line which separates the dark pigmentation of the dorsal and ventral
aspects of the genae.
Material Examined

Arkansas:
     North Sylamore Cr., off Gunner Rd.,
     G. A. Schuster (1 male).
off Ar. 14, Stone Co., Ar., 20-V-74.
Illinois:
     Illinois R., at Starved Rock St. Pk., LaSalle Co., II., 26-VIII-76.  D.
     A. Etnier (3 mmts., many larvae); Kankakee R., at U.S. 45 and 52 jet.
     with II. 115, Kankakee, Kankakee Co., II., 8-IX-76.  D. A. Etnier, G. A.
     Schuster, M. H. Hughes, N. M. Burkhead (many mmts, and larvae); Kankakee
     R., in Momence, Kankakee Co., II., 8-IX-76.  D. A. Etnier, G. A. Schuster,
     M. H. Hughes, N. M. Burkhead (1 male mmt., 12 larvae).

Louisiana:
     Bogue Chitto, at La. 41, St. Tammany Par., La., 8-VI-74.  J. A. Louton
     (many males); Atchafalia R., at Simmsport, Pointe Coupee Par., La., 14-
     VII-75.  J. A. Louton  (13 males); Baton Rouge, E. Baton Rouge Par., La.,
     10-VIII-73.  J. A. Louton (4 males); Trout Cr., at White Sulphur Springs
     on La. 8, LaSalle Par,.,. La., 25-VIII-73.  J. A. Louton (many adults).

Minnesota:                .
     Saganaga Falls, Lake Saganaga, Cook Co., Mn., 3-VIII-75.  D. A. Etnier
     (2 male mmts., several larvae); Mississippi R., at Fort Snelling, Mn.,
     19-VII-75.  D. A. Etnier (1 larva); Saganaga Falls, Lake Saganaga, Cook
     Co., Mn., 22-VII-75.  D. A. Etnier (several mmts. and larvae).
                                      73

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Ohio:
     Ohio R., Beckford Power Plant, 10 mi. upstream from Cincinnati,
     21-VII-75.  WAPORA  (1 larva, several male, and female mints.); Great
     Miami R., Miamitown, Hamilton Co., Oh., 13-VII-75.  D. A. Etnier (6
     male mmts., many larvae); Great Miami R., at Island Park, Dayton,
     Montgomery Co., Oh., 6-VII-76.  G. A. Schuster (2 larvae); Great
     Maimi R., at Island Park, Dayton, Montgomery Co., Oh., 7-VTII-76.
     G. A. Schuster, F. Schuster (1 larva).
Tennessee:
     Reelfoot lake, Tn 22, Obion Co.,
     males, 13 females).

Distribution
Tn., 18-V-74.  G.  A.  Schuster (12
     Published distribution records include the following:  Alabama, Arkansas,
Georgia, Illinois, Indiana, Kentucky, Michigan, Minnesota, Ohio, Texas, and
Wisconsin.  .This general scheme indicates that the distribution of H. orris
lies within the Mississippi and Mobile Basins.

Biology

     This species appears to be associated with big rivers.  It, along with
its sister species, H. bidens, seems to be able to cope with siltation better
than most Hydropsyche species.  Both are often collected in rivers with a
high silt load and high concentration of suspended organic substances.

     H, orris populations are often of staggering density, that is, individual
larval retreats and pupal cases literally stack on top of one another.  This
is probably due to the high concentration of suspended organic matter in the
rivers where they occur.

     Pupal cases are constructed predominantly of secreted substances with
sand grains attached to it.  The pupal case is, therefore, rather flexible
and differs from the general format of Hydropsyche pupal case construction.
Because H. orris populations are so dense, it is not uncommon to find
sloughed larval sclerites incorporated into pupal case construction.  One
case in particularly which was examined had a large number of larval sclerites
embedded within the sand grain structure of the case.  These consisted of
several frontoclypea, nota, labra, and mandibles.  Typical pupal case length
is 8.0 mm; case width is 3.5 mm.

     On several occasions H. orris pupal cases collected in the Saganaga
River at Saganaga Falls, Minnesota, contained Symphitopsyohe recurvata larva
and pupa.  This interesting association has never been noted before.  It
appears that, after H. orris emerges, S. recurvata utilizes^ the empty H.
orris pupal cases in which to pupate.  This allows S. recurvata to conserve
energy that otherwise would have been spent in constructing its own case.
Not all individuals of a S. recurvata population take advantage of this
situation since they are more commonly collected in their own typical rock-
structured cases.
                                      74

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      From published  and  unpublished  collection data,  it  appears that H. orri-s
 is  univoltine,  emerging  from April to  October.

                           Hydropsyche bidens  Ross
                               (Fig.  36A and B)

      Rydropsyche bidens  Ross,  1938a.   111. Nat. Hist.  Surv.,  21:142 (Type
 locality:  Apple River Canyon  St. Pk., 22-VIII-35.  Belong and Ross).

      Eydropsyohe bidens:  Denning, 1943.  Ent. Amer.,  23:109; Ross, 1944.
 111.  Nat.  Hist. Surv., 23:5, 91, 95, 197; Etnier, 1965.  Ent. News, 76:146;
 Unzicker  et al., 1970.   J. Ga. Ent.  Soc., 5:171; Longridge and Hilsenhoff,
 1973.  Wise. Acad. Sci., Arts  and Letters, 61:176.

 Description

      Head  capsule length, 1.30 to 1.50 mm; width, 1.30 to 1.40 mm.  Antero-
 lateral margin  of frontoclypeus with large, conspicuous  teeth.  Color pattern
 of  head with two, large, clear, yellowish, oblique spots in the middle of the
 frontoclypeus.  Lateral margins of these spots contiguous with yellow areas
 around eyes.  Anterior section of frontoclypeus darker than posterior portion.
 Head  without conspicuous black spines, but covered with  small, slender, clear,
 inconspicuous spine-like setae.  Head  in lateral view broadly rounded; flat-
 tened dorsally  except for raised teeth on the frontoclypeus.   Laterally,
 area  around eye yellow.  Directly posterior to eye, a thin, clear line
 separates  dark  coloration of dorsal and ventral aspects  of genae.  Ventral
 aspects of genae with large areas of dark brown pigmentation.  Dorsally, dark,
 coloration along suture line connecting right and left genae.  Thoracic
 sclerites  and legs golden brown to dark brown.  Labrum brown, lateral hair
 fringes similarly colored.  Anterior edge of pronotum almost lacking bristle-
 like  setae hairs.  Pronotum covered with very short spine—like setae that are
 same  as ground color.  Pronotum laterally edged in black as is posterior
 margin.  Pronotum and mesonotum without hairs; metanotum usually has short,
 thin  hairs.  Prosternum with black spots on anterolateral corners and black
 bar on posterior border.  Prosternal plates S-shaped, occasionally separated
 in  center  so that each is divided.  Sternal plates on mesothorax wide and
 slender with lateral margins projecting anteriad.  Ventral gills typical;
 filaments  abundant.  Venter of anal legs covered with numerous,  slender,
 golden spine-like setae.  Scale hairs very dense on dorsum of abdominal
 segments IV - VIII; no scale hairs below pleural gills.  Segments IV -
VIII  also with uniformly spaced, short, black setae.   Abdominal segment III
with  scale hairs spaced further apart, these hairs decreasing greatly on
 segment II, and restricted to a few scale hairs only on lateral aspects of
 segment I.  Abdominal segments I - VIII with .exceedingly small spines  which
 give a peppered appearance under lower magnifications.  Minute spines
 interspersed between scale hairs and black setae which are most prevalent
 on anterior segments, especially on mesal areas of segment I.

Variation

     The material examined showed no distinctive variation either geographi-
cally or within populations.
                                     75

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Diagnosis

     This species is closely related to H. orris.  They are often difficult
to separate in both larval and adult stages.  No consistent morphological
characters could be found to separate the larvae of these two species.
Color patterns of head and ground color of thoracic sclerites and legs
can be used to reliably separate the two species.  The thoracic sclerites
and legs of H. bidens bear golden brown to dark brown coloration; whereas,
on H. orris thoracic sclerites and legs are straw colored.  Differences
in head capsule coloration are more pronounced.  The posterior one-fourth
of the head of H. orris is devoid of coloration, where H. bidens has a
medial posterior dark strip of pigmentation on each side of the suture
connecting right and left genae.  Hydropsyohe bidens typically possesses
two large, clear spots on the frontoclypeus (Fig. 36A).  Hydropsyohe orris
has a large, V-shaped, light spot (Fig. 35A); rarely is it broken into
two spots.  In general, H. bidens has a darker coloration of all sclerites
than H. orris.

Material Examined

Illinois:
     Kankakee R., Momence, Kankakee Co., II., 8-IX-76.  D. A. Etnier,
     G. A. Schuster, M. H. Hughes, N. M. Burkhead (2 larvae); Kankakee
     R., Kankakee, Kankakee Co., II., l-IX-76.  D. A. Etnier, G. A.
     Schuster, M. H. Hughes, N. M. Burkhead (9 larvae, 2 male adults);
     S. Fk. Rock R., at U.S. 51, Rockford, Winnebago Co., II., 25-VIII-
     76.  D. A. Etnier (1 adult male, 3 mints.).

Minnesota                 ;
     CrowR., Watertown, Carver Co., Mn., 17-VII-75.  D. A. Etnier
     (several larvae); Crow R., Watertown, Carver Co., Mn., 27-VIII-75.
     D. A. Etnier  (1 male mint.); Minnesota R., at Montevideo, Chippewa
     Co., Mn., 8-VII-76.  D. A. Etnier  (several mmts. and larvae);
     Chippewa R., at Mn. 15, Montevideo, Chippewa Co., Mn., 8-VII-76.
     D. A. Etnier  (many mmts. and larvae).

Distribution

     Published distribution records include the following states:  Arkansas,
Illinois, Indiana, Iowa, Michigan, Minnesota, Wisconsin, Missouri, Ohio,
and Texas.


Biology

     This species, as in the case of its  sister species, H. orris, is typically
found  in big  rivers.  These two species seem  to be able  to cope with heavy
siltation and suspended materials.  Often it  is found in large numbers with
larval retreats  and pupal cases stacked one on the other.  When  found in
                                      76

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association with H. owls, it is never in the numbers that one finds in E.
orris populations.  It is noteworthy that, when these two species are asso-
ciated, H. bidens is often rare.  This may indicate that H. orris outcompetes
5. bidens.

     Pupal cases are similar to those of H. orris; the case is covered with
sand grains and is flexible.

     Emergence data indicates that H. bidens is univoltine and emerges from
April to September.

                           Hydropsyohe aerata Ross
                                  (Fig. 37)

     Hydropsyche aerata Ross, 1938.  111. Nat. Hist. Surv. Bui., 21:144-145
(Type locality:  Aurora, II., 17-VII-27.  Prison and Glasgow).

     Hydropsyohe aerata:  Ross, 1944.  111. Nat. Hist. Surv. Bui., 23:101-102;
Leonard and Leonard, 1949.  Occ. Pap. Mus. Zoo., Mich., 522:10.

Description

     Head capsule length, 1.45 to 1.60 mm; width, 1.25 to 1.40 mm.  Head yellow
except for dark pattern on dorsum.  Color pattern dark brown and in rough out-
line of a cross with transverse arms wider than longitudinal bar.  Clear spot at
posterior angle of frontoclypeus where transverse and longitudinal bars meet.
Head with scattered short, brown spines, mostly concentrated on dorsum of
genae.  Frontoclypeus with patch of numerous, minute, brown spine—like setae
on midlateral margin.  Labrum brown, covered with long, black setae; laterally
margined with tuft of long, golden setae.  Mandibles yellow; apex dark brown
to black.  Thoracic nota yellow to light brown, covered with short, black
hairs.  A few conspicuous, black bristle-like setae and spine-like setae on
nota.  Legs same color as nota, and adorned with numerous, short, brown spine-
like setae.  Prosternum yellow with black posterior margin.  Poststernal
plates and mesosternal plates brown.  Abdomen beige with numerous, short,
black hairs.  Scale hairs with greatest concentration on dorsum of last three
segments.  Few or no scale hairs on anterior segments, especially segments I -
II.  Interspersed on segments I - IV are numerous, minute spines which give-
peppered appearance under lower magnifications.  Dorsal sclerites of anal legs
with short, black setae; some small, brown spine-like setae laterally.  Venter
of anal legs covered with numerous, short, brown spine-like setae.

Diagnosis                f: "'

     Hydropsyehe aerata is distinguished from other Hydropsy-she species by the
following combination of characters.  First, the color pattern of the head is
unique to the species.  Second, there are no or very few scale hairs on the
first several abdominal segments, the greatest number being on segments V —
VIII.  Last, there are numerous heavily sclerotized brown spine-like setae on
the venter of the anal legs.  Also see comments under diagnosis for H.  frisoni.
                                      77

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 Material Examined

 Illinois:
      Kankakee R., on U.S. 1 S. and 17 W. in Momence, Kankakee Co., II., 8-IX-
      76.  D. A. Etnier, M. H. Hughes, N. M. Burkhead, G. A. Schuster (3 mints.,
      10 larvae); Kankakee R., at U.S. 45 and 52 in Kankakee, Kankakee Co., II.,
      8-IX-76,  M. H. Hughes, D. A. Etnier, N. M. Burkhead, G. A. Schuster (3 :
      mmts.,  2 larvae, 1 adult male).

 Distribution

      The known distribution of H.  aerata includes the following states:
 Illinois, Indiana, and Michigan.

 Biology

      This species is not well known.   Most of the published records of  H.
 aerata are .from the Kankakee River in Illinois.   The two localities,  listed
 above,  are very similar physiographically and in close proximity to one
 another.   The Kankakee River at these localities is a large midwestern  river
 with very wide (100 meters)  and shallow (0.3 to  0.6 meters)  riffle areas.  The
 bottom consists primarily of small-to-medium-sized rocks with a great deal of
 gravel.   The color of the water is brownish with an apparent high suspended
 organic load.   The numbers of net-spinning caddis larvae are very high.   Other
 common Hydropsychids collected at  these localities include:   numerous
 Cheumatopsyche spp.  larvae,  Potamyi-a  flava,  Macronema zebratum,  Hydropsyche
 orris,  H.  bidens,  H.  -Lnoommoda,  H.  simulans,  and H.  phalevata.   All of  these
 can  unquestionably be classified as "big river"  species  since they are  most
 commonly  collected in such habitats.

      Ross (1944)  give May through  August as  the  period of emergence.

                          Eydropsyohe  plialevata Hagen
                                   (Fig.  38)

      Hydropsyche pTmlevata Hagen,  1861.   Syn.  Neur.  N. Am.,  p.  287  (Habitat:
 "St.  Lawrence  R.,  Canada:  Washington (Osten Sacken);  Pennsylvania  (Zimmerman)")

      Hydropsyche phalerata:  Hagen, 1864.  Verh.  Zool. Bot.  Ges., 14:823.
Provancher,  1877.  Nat.  Can.,  9:267;  Provancher,  1878.   Pet.  Faune  Ent. Can.,
2:142; Provancher, 1878. .Nat. Can.,10:147;  Banks,  1892.  Trans; Am. Ent.
Soc., 19:367;  Banks  1894.  Ent.  News,  5:180;  Smith,  1900.  Ins. N.J., p. 64;
Banks, 1904.   Proc.  Ent.  Soc. Wash.,  6:214; Ulmer,  1905.   Zoo.  Insbiol., 1:68;
Ulmer, 1907.   Gen. Ins.,  60:170; Ulmer,  1907.  Cat.  Coll.  Selys  6,  1:66; Banks,
1907.  Cat. Neur.  Ins. U.S., p.  47; Sibley, 1926.   Bui.  Lloyd Libr. 27 Ent.
Ser., 5:104; Betten, 1926.  Mem. Cornell Agric. Exp.  Sta., 101:524; Betten,
1934.  Caddis  Flies N.Y.  St., p. 189;  Banks,  1936..   Psyche,  43:126,  129;
Milne, 1936.   Stud. N. Am. Trich.,  3:73  (as syn.  of moTOsa); Ross,  1938.
Psyche, 45:18; Ross, 1941.  Trans. Am. Ent. Soc.,  67:90; Denning, 1943.  Ent.
Amer., 23:109, 111,  113-114  (Betten 1934 is syn.  of spcama); Ross,  1944.  111.
Nat. Hist. Surv. Bui., 23:15, 86, 91,  93, 102, 294; Leonard and Leonard, 1949.
Occ. Pap. Mus. Zoo., Mich., 522:10; Etnier, 1965.  Ent.  News, 76:146;

                                      78

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 Longridge  and Hilsenhoff,  1973.  Wise. Acad.  Sci., Arts and Letters, 61:176;
 Resh,  1975.  Trans. Ky. Acad.  Sci., 36:12.

 Description

     Head  capsule length,  1.35 to 1.50 mm; width, 1.10 to 1.30 mm.  Background
 color  of head yellow; frontoclypeus with golden brown image of an arrow point-
 ing posteriorad.  Genae with dark brown mottling bordering frontoclypeus.
 Wide transverse band of brown pigmentation with numerous, yellow spots pos-
 terior to  eyes.  Venter of head yellow.  Posterior portion of frontoclypeus
 and dorsolateral surfaces  of genae covered with short, golden spine-like
 setae.  In addition to above setae, genae decorated dorsally and laterally by
 large, conspicuous, black  setae that slightly increase in diameter distally.
 Apical margin of frontoclypeus produced into a low triangular point.  Labrum
 golden brown; covered with thick, black setae and more slender, brown setae;
 laterally  margined with brush of long yellow setae.  Base of mandibles
 yellow; apically dark brown.  Thoracic sclerites slightly darker than head.
 Pronotum and mesonotum covered by large, conspicuous black setae like those
 on head; metanotum with very few such setae.  Nota also covered with short,
 clear, and inconspicuous spine-like setae.  Coxal segments of legs brown;
 all other  segments yellow.  Setation of leg segments similar to that of other
 Hydropsyche species.  Prosternum yellow, posterior margin with wide, black
 band; poststernal plates brown, elongated and pointed laterally.   Mesosternal
 plates triangular.  Abdomen brown.  Dorsum covered with many scale hairs, short,
 black hairs inconspicuous  except on anterior segments.  Venter of abdomen
 lushly covered with short, black hairs but lacking scale hairs.  Exceedingly
 tiny spines cover the dorsum of abdomen, seen clearly only under high magnifi-
 cation.  Greatest concentration of these spines on first several segments.
Dorsal sclerite of anal legs covered with black setae similar to those on
head.  Venter of anal legs with short, black hairs and numerous,  short, clear,
 golden spine-like setae.

Variation

     Larvae have been examined from five different states.   The only variation
 found is darkness of the color pattern of the sclerites of the head and thorax.
It has been observed that those larvae from Minnesota and Illinois have a
greater contrast in color pattern.   On the other hand, the southern material
examined does not show this degree of contrast between the background and the
pattern of the genae and frontoclypeus.   No variation in the anterior margin
of the frontoclypeus has been noted—it always protrudes as a low,  triangular
point.                  •.•'••••-

Diagnosis

     The most diagnostic character permitting the separation of H.  phalerata
from other Hydropsyohe species is the anterior margin of the frontoclypeus
 (Fig.  38).   No other species in the genus has an apical border of the
frontoclypeus which forms a low,  triangular point.
                                      79

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Material Examined

Illinois:
     Kankakee R., in Momence, Kankakee Co., II., on II. 1 S. and 17 W., 8-IX-
     76.  D. A. Etnier, M. Hughes, N. M. Burkhead, G. A. Schuster (several
     larvae); Kankakee R., at U.S. 45 and 52, in Kankakee, Kankakee Co., II.,
     8-IX-76.  D. A. Etnier, M. Hughes, N. M. Burkhead, G. A. Schuster  (1
     larva).

Kentucky:
     Cumberland R., on Cumberland Falls Rd., 5 mi. W. of Ky. 264, Whitley Co.,
     Ky., 29-VI-75.  G. W. Wolfe, G. A. Schuster (several larvae).

Minnesota:
     Mississippi R., at Fort Snelling, Mn., 19-VII-75.  D. A. Etnier  (several
     larvae, 1 male mmt.).

Tennessee:
     Holston R., under 1-40 bridge, Knox Co., Tn., 27-IV-75.  D. A. Etnier (5
     male mints.); Holston R., under 1-40 bridge, Knox Co., Tn., 8-VII-75.  D.
     A. Etnier  (1 male mmt.); Nolichucky R., at Solomon Island of unnamed Co.
     Rd., 12.6 mi. N. of  Newport, Cocke and Greene Co. line, 26-X-75.  D. A.
     Etnier  (several larvae).

Virginia:
     James R., at Va. 45, Goochland Co., Va., 13-VI-75.  D. A. Etnier, W. C.
     Starnes (6 larvae);  James R., at Peters Cr., on Va. 501, Bedford Co., Va.,
     30-VII-76.  D. A. Etnier, G. A. Schuster (several larvae).

Distribution

     Hydropsyche phalerata is widespread over eastern North America.  Published
records  include the following states:  Georgia, Indiana, Illinois, Kansas,
Kentucky, Michigan, Minnesota, New Jersey, New York, North Carolina, Ohio,
Pennsylvania, Tennessee,  Virginia, and Wisconsin.

Biology

     Hydropsyohe phalerata is typically collected in large, warm-water  rivers.
All  the  rivers  in which we have  collected H. phalerata can be classified as:
 (1)  very wide with  long,  shallow riffle areas,  (2) substrate consisting of
small-to-medium-sized rocks with course gravel and often covered with silt,
 (3)  high suspended  organic load,  and  (4) warm water throughout the late spring
and  early  fall months.  Hydropsyohe species often collected with H. phalerata
include  the following:  H. b-idens, H. orris, and H. simulans.  Ross  (1944)
indicates  that H. phalerata emerges from  late April to September.

                          Eydropsydhe dioantha Ross
                                  (Fig. 39)

     Hydropsyche dicantha'Ross,  1938a.  111. Nat. Hist. Surv. Bui., 21:146
(Type locality:  Swansea, Ontario, 15-VIII-34.  H. S. Parish).

                                      80

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     Eydropsyohe d-ioantha:  Ross, 1944.  111. Nat. Hist. Surv. Bui., 23:91,
93, 102, 294; Leonard and Leonard, 1949.  Occ. Pap. Mus. Zool., Mich., 522:9;
Morse and Blickle, 1953.  Ent. News, 64:71; Etnier, 1965.  Ent. News, 76:146;
Longridge and Hilsenhoff, 1973.  Wise. Acad. Sci., Arts and Letters, 61:176;
Resh, 1975.  Trans. Ky. Acad. Sci., 36:12.

Description

     Head capsule length, 1.15 to 1.30 mm; width, 1.00 to 1*10 mm.  Head
brown dorsally and ventrally; yellow laterally, around and behind the eyes.
Frontoclypeus with numerous, stout, black setae, and four distinct yellow
spots.  One pair located medially on sclerite; other two spots located
anterior and slightly lateral to these.  Labrum golden with black setae;
laterally margined with tuft of long, yellow setae.  Mandibles golden boown,
and edged in dark brown.  Thoracic nota golden; laterally trimmed in black.
Pronotum with numerous, short, black setae; mesonotum with less, and metanotum
with still fewer.  Prosternum yellow; anterior margin trimmed in black; black,
bar posteriorly.  Poststernal plates and mesosternal plates brown.  Legs
yellow; first three segments trimmed in black.  All segments amply adorned
with small, brown spine-like setae.  Abdomen beige with numerous, short,
black hairs.  Scale hairs moderately abundant; equally concentrated on all
segments except first, which has much fewer.  Dorsal sclerites of anal legs
golden, covered with short, black setae.  Venter of anal legs covered with
short, black setae, but without large spine-like setae.  Minute spines
present on abdominal segments VIlI — IX.

Variation

     The only variation found in this larva is the shade of the ground color
of the head.  It is sometimes a darker bronze than the typical brown colo-
ration.  This variation has been found to be intra-populational.

Diagnosis

     This species most resembles H* demova and H. valanis to which it is most
closely related.  All three species possess two pairs of distinct, round,
yellow spots on the anterolateral margins of the frontoclypeus.  HydropsyGhe
Valanis with spine-like setae on the venter of the anal legs is easily sepa-
rated from the other two species.  Eydropsyohe d-icantha can be differentiated
from H. demoTa in a number of ways.  First, and probably the most important
diagnostic feature of H. -.demora, is the restriction of scale hairs to the
posterior three segments.  Even on these segments, however, the scale hairs
are very sparse and often difficult to locate without a compound microscope.
Hydropsyche dieantha, on the other hand, has abundant scale hairs on all
segments.  Second, H. d-icantha has numerous, distinct, short, black bristle-
like setae scattered over the entire body of the frontoclypeus.  Hydropsyche
demora., on the other hand, possesses very short, clear spine-like setae on
the posterior half of the frontoclypeus, and anterior half lacks setae.
                                     81

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Material Examined

Kentucky:
     Horselick Cr., off Ky. 89 on unnumbered Co. Rd., Rockcastle Co., Ky.,
     12-V-76.  G. A. Schuster (1 male mmt.); Silver Cr., on Ky. 876, 5 mi.
     W. of Richmond, Madison Co., Ky., 7-VII-76.  C. and G. A. Schuster
     (several mmt., many larvae); Silver Cr., on Ky. 876, 5 mi. W. of Richmond,
     Madison Co., Ky., 13-VI-76.  G. A. Schuster (several larvae).

Minnesota:
     Baptism R., Finland, Lake Co., Mn., 9-VII-76.  D. A. Etnier, M. A. Etnier,
     S. A. Etnier (1 male mmt., 2 larvae).

Ohio:
     Stillwater R., Deweese  Pk., Dayton, Montgomery Co., Oh., 6-VII-76.  G.
     A. Schuster (3 larvae).

Distribution

     Hydropsy che dicantha appears to have a northerly distribution, restricted
to the eastern U.S.  Published state and province records include:  Kentucky,
Michigan, Minnesota, New Hampshire, New York, Ontario, Washington, D. C., and
Wisconsin.

Biology

     Little is known of the biology of H. diaantha; however,  some insight may
be obtained by the extremely variable habitats  in which  it has been found.
The  four localities listed above represent an extremely  wide  range of environ-
mental conditions.  Horselick Creek, for example, has a  substrate consisting
primarily of sand and mud with few small rocks  in the riffle  area, while Silver
Creek consists chiefly of algae-covered bedrock and  few  small, loose rocks.
Both, however, are rather small, warm-water  streams,  6   to 12 meters wide, and
extremely shallow in the riffle areas  (0.2 to 0.5 meters deep).  The Stillwater
River in Dayton, Ohio, can be classified as  a large  river with heavy
siltation and a  substrate consisting of small-to-large rocks.  On the other
hand, the Baptism River locality represents  a medium-sized trout stream with
fairly cold water throughout the year.

     From the above information one can draw the conclusion that H. dicantha
does not have rigid environmental requirements.  It  also follows that H.
dicantha should  be much more common than it  apparently is.  It is indeed
widespread, but  is by no means common  throughout its range.   This would
indicate that other factors prevent this species from becoming more common.
Two  of these may be predation and interspecific competition.  However,  until
this species is  better understood,  these remain only plausible conjectures.

     Published  collection records indicate  that this species  emerges  from
late June through  September.
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                            Hy dropsy ehe demora Ross
                                   (Fig. 40)

      Hydropsyche demora Ross, 1941a.   Trans.  Amer. Ent.  Soc.,  67:86-87 (Type
 locality:   Demorest,  Ga.,  l-VII-39.  P. W.  Fattig).

      Hydropsyche demora:   Ross,  1944.  111. Nat.  Hist.  Surv. Bui.,  23:294.

 Description

      Head  capsule length,  1.40 to  1.60 mm; head capsule  width,  1.25 to 1.40
 mm.   Frontoclypeus reddish brown with four distinct spots  on anterior  half of
 sclerite;  one pair mesally located with second pair anterolateral to these.
 Posterior  half of frontoclypeus  with  numerous,  small, yellow pits containing
 short,  clear spine-like setae; often  many of  these missing, leaving empty
 pits.   Also,  on posterior  half of  sclerite are 9-10 large, yellow,  muscle
 scars.   Genae brown bordering frontoclypeus;  large,  brown  area  behind  eyes,
 and  large,  yellow areas lateral  to gular suture.   Brown  areas inundated with
 numerous,  yellow pits with short,  clear spine-like setae.  Genae with  scat-
 tered,  long,  black setae dorsolaterally.  Numerous,  large, yellow muscle scars
 on dorsal  mesal border  of  genae; from middle  of frontoclypeus posterior.
 Yellow  muscle scars on  large,  brown areas behind  eye in  two to  three concen-
 tric semicircular rows.  Lateral aspects of head  around  and behind  eyes yellow.
 Venter  of  head brown, especially on stridulatory  surfaces; with numerous,
 yellow,  oblong muscle scars,  particularly posteriorly.   Labrum  golden, clear
 laterally with short, dense,  yellow setae, more abundant, long, brown  setae,
 and  laterally margined  with tuft of long, yellow  setae.  Thoracic nota golden,
 pronotum slightly darker.  Numerous,  short, clear spine-like setae  on prono-.
 turn; mesonotum with fewer; metanotum with none.   Scattered, long, stiff, black
 setae on pronotum;  mesonotum with  several such  setae; metanotum with none.
Mesonotum and  metanotum with  short, thin, black hairs'; pronotum with few or
none.  Pronotum with numerous, yellow muscle  scars laterally;  fewer near
posterdorsal margin.  Prosternum yellow; black  posteriorly.  Poststernal
plates and mesosternal  plates brown.  Legs same color as nota,   and abundantly
adorned with  short, brown spine-like setae.   Abdomen brown with short,  black
hairs dorsally  and ventrally.  Scale hairs,  restricted to last   three segments,
inconspicuous and very  sparse.  Dorsum of segments I - VII with numerous,
minute, brown  spines; becoming smaller posteriorly.  Dorsal sclerite of anal
legs yellow, covered with black setae.  Venter of anal legs with numerous,
short, brown hairs.

Variation                ,                     •

     The small amount of variation observed in this species is  restricted to
the shade of the color pattern on the top of the head.   In some larvae  the
pattern is dark reddish brown, while in other individuals,  it  is more golden.

Diagnosis

     This species may be, at first glance,  confused with H.  dicantha to which
it is closely related.   Both species possess  the four distinct  spots on the
frontoclypeus, but they differ in a number  of  other ways.  First,  and probably

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the most important diagnostic feature of H. demora, is the restriction of
scale hairs to the posterior three segments.  However, even on these three
segments, the scale hairs are,very sparse and often difficult to locate
without a compound microscope.  H. d-lean-tha, on the other hand, has abundant
scale hairs on all except the first two segments.  Second, H. demora possesses
very short, clear spine-like setae on the posterior half of the frontoclypeus.
Eydropsyche dioant'ha has numerous, distinct, short, black setae scattered over
the entire body of the frontoclypeus.  These are absent in H. demora.  Third,
H. demora possesses numerous, yellow muscle scars on the frontoclypeus, dorsum,
and venter of the genae; these are lacking in H. dicantha.  Hydropsyche valanis
also possesses the four yellow spots on the frontoclypeus and may look somewhat
like H. demora.  These two species can be separated by the presence of spine-
like setae on the venter of the anal legs of E. Valanis and their absence on
H. demora.

Material Examined

Tennessee:
     Conasauga R., at Tn. 74 bridge, Bradley Co., Tn., 4-V-75.  C. A. Schuster,
     G. A. Schuster, D. A. Etnier, and Etnier family  (numerous larvae); Little
     R., at Alcoa Water Treatment Plant, 0.5 mi. E. of Tn. 33, Blount Co., Tn.,
     22-VII-75.  G. A. Schuster (2 malemmts., several larvae).

Distribution

     Hydropsyohe demora was previously known only from the type material.  The
known distribution of this species now includes Georgia and eastern Tennessee.

Biology

     Very little is known of the biology of this species.  The two localities
from which larvae and/or metamorphotypes have been collected are quite
different.  That section of the Little River is approximately 4.8 kilometers
upstream from the impoundment of Fort Loudon Reservoir.  The river here is
wide and divided by an island.  The larvae and pupae were all collected on
the south side of the island where there is a 5.0 meter-wide riffle approxi-
mately 0.5 meters deep with very  strong current.  The substrate consists of
medium-size rocks and large broken pieces of concrete from an old bridge
abutment.  The water is brown and heavily laden with  suspended materials, both
organic and inorganic; this causes a great amount of  siltation.  Other Hydro-
psyche species collected here include H. betteni, H. phalerata, and H.
venularis.

     The Conasauga River locality is in direct contrast with the above col-
lecting site.  It is a beautiful  stream with lush aquatic vegetation growing
on a predominantly sand and gravel substrate, with small-to-medium-size rocks
in the riffle area.  The water is extremely clear with little observable
siltation.

     The only feature characteristic to both of  these localities  is  that both
directly drain mountainous terrain.  It is  quite possible that the Little
River locality, before  the impoundment of Fort Loudon Lake, was very similar

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 to the Conasauga River and that H.  demoTa has been able to survive despite
 environmental changes.  It should also be mentioned that the two rivers are
 in completely different drainage basins.   The Conasauga River is in the Mobile
 Basin, and the Little River is in the Tennessee River drainage.

                           Hy'dropsy'che vdlan-Ls Ross
                                   (Fig. 41)

      Hydropsyche valan-is Ross, 1938.   111. Nat. Hist. Surv.  Bui., 21:144
 (Type locality:   Rockton, II., 2-VII-31,  along Rock River.  Prison, Betten,
 and Ross).

      Hydropsy che vdlani-s:  Ross, 1944.  111.  Nat.  Hist. Surv. Bui., 23:6,  91,
 95, 105,  294; Etnier,  1965.  Ent. News, 76:146; Resh, 1975.   Trans. Ky. Acad.
 Sci., 36:12.

 Description

      Head  capsule length, 1.25 to 1.35 mm;  head capsule width, 1.05 to  1.15 mm.
 Posterior  one-fifth of top of head  yellow;  rest of head brown.   Frontoclypeus
 with two pairs of distinct,  round,  yellow spots on the anterolateral portions
 of sclerite.  Posterior half of frontoclypeus with a number  of small, yellow
 spots.  Genae yellow ventrally and  around eye;  margins bordering epicranial
 arms brown which posterior to eye widen and come to level  of eye.   Genae on
 sides and  top with numerous.,  bristle-like, black setae.  Frontoclypeus  with
 similar setae scattered on posterior  half of  sclerite.   Labrum dark brown with
 black setae;  laterally margined with  long, yellow  setae.   Mandibles yellow
 with black margins.  Mesonotum and  metanotum  yellow;  pronotum brown.  Each
 notum with numerous, short,  black,  bristle-like setae.   Prosternum yellow and
 bordered anteriorly and posteriorly in black.   Prosternal  and mesosternal
 plates brown  and not heavily sclerotized.  Legs yellow;  anterior pair slightly
 darker.  All  legs abundantly adorned  with spine-like setae and longer black
 setae.  Abdomen  tan colored;  covered  with numerous scale hairs not noticeably
 decreasing in size or  number anteriorly.  Minute spines  on dorsum of all abdo-
 minal segments.   Scale hairs on sides of  abdominal segments  below pleural gills.
 Venter of  anal legs with numerous sclerotized  spine-like setae;  these setae
 heavily sclerotized at  base;  and long and slender  apically.   Setae on sclerites
 of  last two abdominal  segments  similar  to  those on anal  legs.

 Diagnosis

      This  species  most  resembles E, demova and' E.  d-icctntha,  to which it  is  most
 closely related.   All  three  species possess two pairs of distinct,  round, yel-
 low spots  on  the  anterior  lateral margins of the frontoclypeus.  Bydropsyahe
 vaian-is is most  easily  separated from E. demova and E. dicantha  on the basis -
 of  the spine-like  setae  on the '<• venter of  the anal  legs.  Hydropsy che valanis
 possesses  these  setae, but they  are absent on the  other two  species.  Bydro-
psycl%e •oalanLs may be  distinguished from other  species with  such setae on the
 anal  legs by  a combination of the following characters:  (1)  color pattern  on
 the head,  (2) scale hairs below  the pleural gills  on the abdomen,  (3) bristle-
 like  setae on the  posterior half of the frontoclypeus,  (4) spine-like setae on
 anal  legs are long and rather slender while they are shorter  and more heavily

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sclerotized in the other species.

Material Examined
Ohio:
     Greater Miami R., at Miamitown, Hamilton Co., Oh., 13-VII-75.  D. A.
     Etnier (7 larvae); Greater Miami R., at Island Park, Dayton, Montgomery
     Co., Oh., 6-VII-76.  G. A. Schuster (2 male mmts., 1 larva); Greater
     Miami R., at Island Park, Dayton, Montgomery Co., Oh., 7-VIII-76.
     G. Schuster, F. Schuster  (2 larvae).

Distribution

     Published records include the following states:  Illinois, Indiana, Iowa,
Kentucky, Minnesota, and Wisconsin.  Additional to this list are the above
Ohio records.

Biology

     The biology of  this species is not well understood even though it is
fairly widely distributed.  Ross (1944) indicates that H. valanis may prefer
larger rivers.  The  Greater Miami River, from which the known larvae and
metamorphotypes were collected, is a large, warm-water river.  The river is
approximately 60 to  100 meters wide with very large riffle areas.  The bottom
consists of various-sized, limestone rocks over sand  and  coarse gravel.  It
has a high silt load with a great deal of organic suspended material.

     Species collected with H. volants include H. orris,  B. dicantha, and ^
Symphitopsyche cheilon-Ls.  Symphitopsyche ahe-Llonis is the predominant cad'dis-
fly in the river and is present in astounding numbers.  Hy dropsyche valanis
appears  to be much rarer and  is not commonly encountered.

     Ross  (1944) indicates that emergence of H. valan-is takes place from May
to late  August.

                          Hydropsy ahe ap-indle Ross
                                   (Fig.  42)

     Hydropsyche connate Ross, 1938a.  111. Nat.  Hist. Surv. Bui.,  21:144
 (Type  locality:  Oregon, Illinois,  18-VII-27.   Prison and Glasgow).

     Hydropsyehe arinalei  Ross,  1944.   111. Nat.. Hist.  Surv. Bui.,  23:6, 87,
 91,  93,  104,  294; Unzicker  et al.,  1970.   J. Ga. :Ent. Soc.,  5:171;  Longridge
 and Hilsenhoff,  1973.   Wise.  Acad.  Sci., Arts  and Letters,  61:176.

 Description                                  • ' "'

     Head capsule length,  1.35 to  1.50  mm;  head capsule  width,  1.20 to  1.35
 mm.  Head mostly brown with yellow areas around eyes  and at posterolateral
 sections of genae.   Frontoclypeus  unicolored;  no distinct light spots.   Venter
 of head lighter than top,  stridulatory surfaces slightly darker.   Dorsolateral
 aspects of genae with short,  brown, bristle-like setae;  bristle-like setae

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 present only on  anterolateral  corners  of  frontoclypeus.   Posterior half  of
 frontoclypeus with numerous, thin,  clear, silky setae; present  also on genae
 and all nota.  Bristle-like setae lacking on nota.  Labrum brown with  numerous,
 long, brown setae; laterally fringed by long brush  of yellow setae;  mandibles
 brown.  Nota and legs brown; legs covered with  numerous,  long,  black setae and
 short, brown spine-like  setae.  Abdomen brown;  sparsely covered by normal
 setae and scale  hairs.   Scale  hairs small, 1/2  - 3/4 length  of  regular appres-
 sed hairs.  Minute spines present on dorsum  of  all  abdominal segments  and ven-
 ter of last two  abdominal segments  and anal  legs.   Large,  heavily  sclerotized
 spine-like setae lacking on venter  of  anal legs.

 Diagnosis

     The larvae  of H. apinale  can be distinguished  by a combination of char-
 acters.  These characters include:  (1) the  lack of a distinct  color pattern
 on the head and  the presence of a unicolored frontoclypeus,  (2)  the presence
 of many silky, clear, thin setae  on the posterior half of  the frontoclypeus,
 (3) the lack of  bristle-like setae on  all three thoracic sclerites,  (4)  the
 sparseness of scale hairs on the  abdomen, and (5) the scale  hairs  are  1/2 -
 3/4 the length of regular appressed hairs on the  dorsum of the  abdomen.

Material Examined

 Illinois:
     Indian Cr., Baker,  II., 12-V-38.   Ross  and Burks (1 male,  1 female mint.);
     Indian Cr., Serena, II., 16-V-38.  Ross  and  Burks (2 larvae).

Oklahoma:
     Honey Cr.,  T.F.S.P. Ok., 29-IV-39.  J.  A. and H. H.  Ross (2 male mmts.,
     6 larvae).

Distribution

     Published records of H.  ca"i.nale include the  following states:  Arkansas,
Kansas, Missouri, Oklahoma,  and Wisconsin.  Ross  (1944) describes the range
of H.  arinale as seeming "to follow rather closely the outer  fringe of the
oak-hickory forest."

Biology

     Ross (1944)  states:  "It shows a preference for such streams as Indian
Creek, which is relatively clear and provided with many riffles,or rapids.
Our only large collections, of -adults were taken in May, but emergence con-
tinues through August."    '. .r,

                         Hycbeopsyche soalaris Hagen
                                  (Fig. 43)

     Hydpopsyche SGolaP-Ls Hagen, 1861.   Syn.  Neur. N.  Am., p. 286-287
("Habitat:   St. Lawrence R.,  Can.  (Osten Sacken); Washington; N. Red R.
(Kennicott)").
                                     87

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     Eydropsyche scalaris:  Hagen, 1864.  Verb.. Zool. Bot. Gas., 14:823;
Packard, 1876.  Guide Study Ins. Ed., 5:621; Banks, 1892.  Trans. Am. Ent.
Soc., 19:367; Banks, 1892,  Ent. News, 5:180; Kellogg, 1895.  Am. Natur., 29:
550; Smith, 1900.  Ins. N.J., p. 64; Betten, 1901.  Aquat. Ins. Adirondacks,
p. 573; Banks, 1904.  Proc. Ent. Soc. Wash.., 6:214; Banks, 1904.  Trans. Amer.
Soc., 30:109; Banks, 1905.  Trans. Amer. Ent. Soc., 32:14; Kellogg, 1905.
Amer. Ins., fig. 334; Ulmer, 1905.  Zool. Insbiol., 1:68; Ulmer, 1907.  Gen.
Ins,, 60:171; Ulmer, 1907.  Cat. Coll. Selys., 6, 1:65-66; Banks, 1907.  Cat.
Neur. Ins. U.S., p. 47; Banks, 1908.  Proc. Ent. Soc. Wash., 9:155; Sleight,
1913.  J. N.Y. Ent. Soc., 21:6; Dodds and Hisaw, 1925.  Ecology, 6:386; Essig,
1926.  Ins. W. N. Am., p.;177; Betten, 1926.  Mem. Cornell Ag. Exp. Stat.,
101:524; Muttkowski, 1929*.  Roosevelt Wildlife Ann., 2:192; Betten, 1934.
Caddis Flies N.Y. St., p. 190-191; Neave, 1934.  Int. Rev. Hydrobiol., 31:169;
Milne, 1936.  Stud. N. Am. Trich., 3:69, 72, 73; Banks, 1936.  Psyche, 43:127,
129; Brimley, 1938.  Ins. N. Car., p. 252; Balduf, 1939.  Bion. Entomorph. Ins.,
2; Denning, 1943.  Ent. Am., 23:109, 112-113; Ross, 1944.  111. Nat. Hist.
Surv. Bui., 23:91, 95, 106, 294; Etnier, 1965.  Ent. News, 76:146; Edwards,
1966.  J. Tenn. Acad. Sci., 41:120; Nimmo, 1966.  Can. Ent., 98:691; Corbet
et al., 1966.  Can. Ent., 98:1291; Blicke and Morse, 1966.  Me. Ag. Exp. Stat.
Tech. Bui., 24:6; Unzicker et al., 1970.  J. Ga. Ent. Soc., 5:171; Longridge
and Hilsenhoff, 1973.  Wise. Acad. Sci., Arts and Letters, 522:176.

Description

     Head capsule length of northern larvae, 1.95 to 2.30 mm; head capsule
width, 1.70 to 1.95 mm.  Head capsule length of southern larvae, 1.50 to 1.70
mm; head capsule width, 1,30 to 1.45 mm.  Head reddish yellow laterally,
ventrally and posterodorsally.  Frontoclypeus irregularly mottled brown
against reddish yellow or yellow background.  Genae with brown pigmentation
bordering suture with frontoclypeus and each other. Posterior to eyes genae
also with wide, mottled, brown mark joined to pigmentation bordering suture
of frontoclypeus and genae.  Ventral to this mark slightly ventrally bowed
row of seven to nine circular,   yellow muscle scars outlined in brown.  Below
this two additional rows of muscle scars not as distinctly marked.  Posterior
half of frontoclypeus with numerous, scattered, small, brown spine-like setae
positioned in dark brown pockets.  Dorsum of genae with similar setae, and
scattered long, stiff, bristle-like, black setae.  Some shorter, less stiff
setae located behind and below eyes.  Labrum brown with long, brown setae;
laterally margined with tuft of long golden hairs.  Mandibles brown, trimmed
with dark brown to black.  Thoracic nota brown; pronotum slightly darker than
others, and with numerous, round, yellow muscle scars laterally.  Long, black,
stiff setae on pronotum; less in number on mesonotum; fewer still on metanotum.
Numerous, short, golden spine-like setae on prohbtum and mesonotum;  few to
none on metanotum,-  Conspicuous, short, thin, black hairs cover metanotum;
fewer present on mesonotum; very few on pronotum.  Legs brown; anterior legs
slightly darker.  All segments abundantly garnished with short, brown spine-
like setae.  Abdomen brown with numerous, short, fine hairs.  With exception
of first segment, scale hairs as abundant on dorsum of abdominal segments as
short, fine hairs.  Scale hairs on segment I very  few in number.  Dorsal
sclerite of anal legs with fine, black setae; venter covered with numerous,
short, heavy, brown spine-like setae.
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Variation
     The greatest variation observed immediately upon studying the larvae is
the distinctive size differences between northern and southern larvae.  At
first, it was assumed that these were different species, but upon closer
scrutiny, this was questioned.  On the basis of adult male genitalia, the two
could not be separated, and, other than differences in size, the larvae also
seemed identical.  It is still quite possible that these two forms are sub-
species or possibly valid species.  On the presumption that the larvae are
identical except in size, they are treated here as one species.

     The only other variation either within or between populations is the
ground color pigmentation of the sclerites.  This varies from yellow or
straw colored to a reddish yellow or brown.

Diagnosis

     The following combination of characters may be used to identify H.
sodlaris larvae.  First, the presence of the minute, brown spines on the
dorsum of all abdominal segments.  Second, small, but distinct, spine—like
setae are present on the posterior half of the frontoclypeus, the entire pro-
notum, and the mesonotum where they are fewer in numbers.  Third, there are
numerous, heavily sclerotized spine-like setae on the venter of the anal legs.
Fourth, the color pattern of the head is distinctive, especially the several
slightly curved rows of yellow muscle scars behind the eyes.

Material Examined

Minnesota:
     Mouth of Sunrise R., Chisago Co., Ma., 20-VII-75.  D. A. Etnier (1 male
     mmt., 4 larvae); Sunrise R., at Mn. 95, Chisago Co., Mn., 6-VII-76.  D.
     A. Etnier (several mmts., several larvae).

Tennessee:
     Bull Run Cr., on Tn. 62, just S. of Oak Ridge, Knox Co., Tn., l-IV-76.
     D. A. Etnier, G. A. Schuster (several mmts., many larvae).  Bull Run Cr.,
     on Tn. 62, just S. of Oak Ridge, Knox Co., Tn., 18-IV-76.  D. A. Etnier
     (3 male mmts.).

Virginia:
     James R., at Peters Cr., on Va. 501, Bedford Co., Va., 30-VIII-76.   D. A.
     Etnier, G. A. Schuster (1 male, 1 female mmt., and 5 larvae); New R.,
     below U.S. 460 bridge,,Giles Co., Va., 29-V-75.  G. A. Schuster, D. A.
     Etnier (several larvae).

Wisconsin:
     Red Cedar R., at Co. Rds. M and•W, 22 Mile Ford, Dunn Co., Wi., 3-VII-76.
     D. A. Etnier (several mmts., several larvae).

Distribution                        ' '  '

     In the literature it is reported to be one of the most widely recorded

                                     89

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species in the genus.  Ross  (1944:106) states:  "The name soa1ca"ls is the one
under which many species have been confused.  The selection of a lectotype by
Banks (1936b, p. 172) has given us a definite concept of this species for the
first time."  It is more than likely that the range indicated by earlier pub-
lished records was somewhat  exaggerated from its true range due to this con-
fusion.  Ross (1944) lists the following:  Georgia, Indiana, Missouri, Okla-
homa, Ontario, and Wisconsin.  More recently published records include:
Arkansas, Maine, Minnesota,  Quebec, and Tennessee.

Biology

     Even though this species was often reported in the literature, nothing
was previously known of its  larval biology.  Regardless of the fact that it
has been associated, still little is' known of H. soalaris.  The localities
from which larvae and metamorphotypes were collected are rather different.
They range from a small stream such as Bull Run Creek, with small rocks and
gravel, to the New River, which is a large river with a great deal of bedrock.
However, all the localities  listed above have one thing in common; they can be
termed warm-water,  smallmouth bass-type streams.

                          Hydpopsyche s'imulans Ross
                   (Fig. 2,  3, 4, 5, 6, 9, 10, 44A and B)

     Hydropsyche s-mulons Ross, 1938.  111. Nat. Hist. Surv. Bui., 21:139-141
(Type locality:   Mount Carmel, II., September 11, 1937, along Wabash River.
H. H. Ross).

     Hydeopsyche simulans:  Denning, 1943.  Ent. Am., 23:109, 111, 117-118;
Ross, 1944.  111. Nat. Hist. Surv. Bui., 23:5, 13, 87, 91, 95, 104-105, 294;
Etnier, 1965.  Ent. News, 76:146; Edwards, 1966.  J. Tenn. Acad.  Sci., 41:
120-121; Unzicker et al., 1970.  J. Ga. Ent. Soc., 5:172; Longridge and
Hilsenhoff, 1973.  Wise. Acad. Sci., Arts and Letters, 522:176;  Edwards, 1973.
Tx. J. Sci., 24:504; Resh, 1975.  Trans. Ky. Acad. Sci., 36:12.

Description

     Head capsule length, 1.65 to 1.85 mm; width, 1.50 to 1.70 mm.  Dorsum of
head dark brown with pair of large, yellow spots on frontoclypeus.  Laterally,
genae with a large irregular yellow area, which posteriorly reaches dorsum of
head.  Venter mostly dark brown, especially on stridulatory surfaces and
surrounding gular suture.  Dorsolaterally genae with numerous, stout, black
setae, and small, golden spine—like setae.  Posterior to frontoclypeus, genae
with numerous, oblong, yellow muscle scars on dark brown background.  Labrum
straw colored with long, black setae and laterally margined with tuft of long,
yellow satae.  Mandibles straw colored, dark brown apically.  Pronotum brown,
other nota straw colored.  Each notum with scattered, stout, black setae;
short, black hairs; and short, golden spine-like setae.  Prosternum, post-
sternal plates,  and mesothoracic plates yellow; prosternum with posterior
black bar.  Legs straw colored to yellowish brown and covered with numerous,
golden spine-like setae.  Abdomen light brown covered with short, black setae
both dorsally and ventrally.  Dorsum also with numerous scale hairs most
numerous on posterior segments.  Interspersed between scale hairs and other

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 hairs are minute golden spines that are a peppered appearance under lower
 magnifications.  Spines most numerous on segments I - IV; on more posterior
 segments, these spines extremely minute, less abundant, and distinctly visible
 only under oil emersion.  Anal legs covered dorsally and ventrally with short
 black hairs.

 Variation

      There is limited variation of color pattern within and between populations
 of H.  svrmlans.  The large, yellow areas on the side of the head may be inun-
 dated with brown pigment but never to the degree of H.  hagen-i or H.  leonardi.
 The only variation in the dorsal coloration is a difference in the size of the
 spots on the frontoclypeus.  They may sometimes be larger and more conspicuous
 than in typical specimens (Fig.  3).

 Diagnosis

      This species is best separated  from other soalaris group species by the
 color pattern of the head.   The  only species  with which it may be confused are
 H.  hageni or H.  Zeonardi.   These two species  also possess a pair of  spots on
 the frontoclypeus which are not  as large or conspicuous as those of  H.  s-mulans.
 Hydropsyohe  hageni and H.  leonard-L also  possess a much  darker background color,
 being  very dark to almost black.   Hydropsyche simulans  also possesses numerous,
 yellow muscle scars on the posteromesal  sections of  the genae.

 Material Examined

 Illinois:
     Kankakee R.,  on U.S.  1 S. and 17 W.  in Momence, Kankakee Co., II.,  8-IX-
     76.   D.  A.  Etnier, M.  H. Hughes, N. M. Burkhead, G.  A.  Schuster  (several
     mmts. and larvae); Kankakee R., at  U.S.  45 and  52  in Kankakee, Kankakee
     Co.,  II.,  8-IX-76.  M.  H. Hughes, D. A.  Etnier, N. M.  Burkhead,  G.  A.
     Schuster (3 mmts.).

 Kentucky:
     Cumberland R.,  on  Cumberland  Falls  Rd.,  5  mi. W. of Ky.  264, Whitley Co.,
     Ky.,  29-VI-75.   G. W. Wolfe,  G. A.  Schuster  (1 male mmt., several larvae).

Minnesota:
     Chippewa R., at Mn. 15, Montevideo, Chippewa Co., Mn., 8-VII-76.  D. A.
     Etnier  (3 mmts., several larvae).

Tennessee:                   ---_
     French Broad R., one-fourth mi.  below Johnson Bible College, Knox Co.,
     Tn., 27-VI-75.  D. A. Etnier, G. A. Schuster (several mmts. and many
     larvae); Holston R., under 1-40 bridge, Knox Co., Tn., 8-VII-75.  D. A.
     Etnier  (1 male mmt., several larvae).

Texas:
     Brazos R., Hwy. 4 bridge, Palo Pinto Co., Tx., 24-11-76.  K. Stewart
     (many larvae and mmts.).
                                      91

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Distribution

     Eydropsydhe simulans is widespread over the central U.S. and also occurs
in some of the more easterly states.  State records of this species include
the following:  Illinois, Indiana, Iowa, Kansas, Kentucky, Minnesota, Missouri,
Ohio, Oklahoma, Tennessee, Texas, and Wisconsin.

Biology

     Larvae of S. simulcms have only been collected from large rivers with
long, wide riffle areas.  The Cumberland, Holston, and French Broad localities
listed above basically represent similar habitats.  In each case, the rivers
are very wide  (30 to 60 meters), fairly muddy, and with large riffle areas.
The substrate consists typically of medium-to-large-size rocks with abundant
coarse gravel.  Eydvopsyohe simulans seems to prefer the larger rocks and
avoids the smaller ones which tend to be washed downstream by the current.
The larval retreats and pupal cases are most often found on or near the bottom
of the rocks with cracks and crevices used as anchoring points.  Perhaps due
to the large amount of organic material in the water, E. simulans is usually^
found in large numbers.  Other Eydropsyche commonly collected with this species
are E. phalerata and E. owls.

     Ross (1944) indicates emergence of this species takes place from April to
September.

                         Eydropsyche inoammoda Hagen

     Eydropsyche ineammoda Hagen, 1861.  Syn. Neur. N. Am.,  p. 290-291  (Type
locality:  "Georgia  (Collection of Hagen)").

     Eydropsyche insormodai  Hagen, 1864.  Verh.  Zool. Bot.  Ges., 14:822;
Banks, 1892.   Trans. Amer. Ent. Soc., 19:367; Ulmer 1905.  Z.  Insbiol., 1:68;
Ulmer, 1907a.   Gen.  Ins., 60:171; Ulmer, 1907b.   Cat.  Coll.  Selys.,  1:68;
Banks, 1907.   Cat. Neur.  Ins. U.S., p.  47; Krafka, 1923.  J. N.Y. Ent.  Soc.,
31:39, 45; Krafka, 1924.  Ann. Ent. Soc. Amer.,  17; Betten,  1934.  Caddis
Flies N.Y. St.,  p. 188  (not  sensu Hagen); Betten,  1936.   Mem.  Cornell Ag.
Exp. Stat.,  101:524; Milne,  1936.   Stud. N. Am.  Trich.,  3:73 (as syn. of
sealaris'); Banks,  1936.   Psyche,  43:128, 129  (distinct species); Ross,  1938.
Psyche,  45:17; Ross,  1944.   111.  Nat. Hist. Surv.  Bui.,  23:15,  91, 95,  106,
294; Unzicker et al.,  1970.  J.  Ga. Ent. Soc.,  5:171;  Resh,  1975.  Trans.  Ky.
Acad.  Sci.,  36:12.

Diagnosis

     Eydropsyohe inoormoda  larva cannot at  this,time  be distinguished from E.
simulans (see Fig.  3).   Larvae and  pupae have only been collected  from the
Kankakee River at  Kankakee,  Illinois.   Perhaps with the collection of more
material of  this widespread species,  it will_be better understood, and char-
 acters can be found to separate it  from E.  simulans.
                                       92

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Material Examined

Illinois:
     Kankakee R., at U.S. 45 and 52, Kankakee, Kankakee Co., II., 8-IX-76.
     M. H. Hughes, D. A. Etnier, N. M. Burkhead, G. A. Schuster  (1 male,
     3 female mmts,, 5 larvae).

Distribution

     This species has been recorded from Arkansas, Florida, Georgia, Illinois,
Kentucky, Louisiana, New York, and North Carolina.

Biology

     Even though this species is widespread in eastern North America, little
is known of its biology.  The Kankakee is a larger river with wide riffle
areas in which H. ineammoda was collected.  It was collected with H. orris,
ff, phalevcrta, H. b-uL&ns, and H. aerata.  Ross (1944) indicates that H.
-£rb3
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these present but scattered and inconspicuous on posterior half of fronto-
clypeus.  Longer brown setae along sutures between genae and frontoclypeus.
Labrum yellow with brown setae, fringed laterally with longer yellow setae.
Mandibles with yellow base and dark brown apices.  Thoracic nota yellow.
Pronotum with anterior border darkened; entire notum covered with scattered
short golden spine-like setae and dark brown hairs.  Mesonotum also covered
with golden, spine-like setae and longer, darker hairs than those of pronotum;
and with a "U-shaped" black mark posteriad.  Metanotum lacking spine-like setae
and with still longer, darker hairs; arid with a median posterior black spot.
Legs yellow with reddish brown claws, scattered long black setae, and covered
with short, golden, spine-like setae.  Abdomen with minute spines on first
five segments.  Scale hairs sparse on first two segments, gradually becoming
slightly more dense on posterior segments, and absent below pleural gills.
Bases of anal legs with thickened golden qpine-like setae basally and
ventrolaterally.

Diagnosis

     The pale yellow color of the head capsule and thoracic nota are shared
only with H. scalaris, H. patera* H. phalerata, and H. aerata.  In both
sealarfis and patera the pronotum is consistently darker than the other
thoracic' nota, whereas in 'frison-i- the nota are equally pale except for a
slightly darker anterior border on the pronotum.  Differs from phalerata in
having the anterior border of the frontoclypeus straight.  Most similar to
ff. aevata from which it differs in having less conspicuous spine-like setae
on the ventral bases of the anal legs.  In aerata these setae are of approxi-
mately the same size, shape, and color as those on the sclerites on the ven-
tral surface of the ninth segment; in fri-sorri these setae are much smaller
and less robust than those on the sclerites on the venter of the ninth
segment,  H. frtsoni lacks the brown blotch surrounding the epicranial stem
that is present in aerata, and has the posterior angle of the frontoclypeus
brown rather than with a pale spot as in aerata.  In H. aerata scale hairs
on the abdomen are much more abundant on posterior than on anterior abdominal
segments, whereas in fri-sori'C there is a less noticeable increase, in their
abundance from anterior to posterior.

Variation                  '

     In addition to slight variation noted in the extent of brown pigment on
the frontoclypeus (see Description), there is occasionally a small brown
blotch in the anterolateral corner of the frontoclypeus.

Material Examined                              ._. „ •",."

Alabama:
     Cahaba River at Co. Hwy. 27 bridge, Bibb',Co...'," Ala., 15-IV-78.  B. H.
Bauer, D. A. Etnier (many larvae); Little Cahaba River at Bulldog Bend,
along co. rd. 65, T24N, R 11E, S 25, Bibb Co.", Ala., 15-IV-78.  B. H. Bauer,
D. A. Etnier (1 male from spider webb, 3 male mmts., many larvae).
                                     94

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Minnesota:
     Mouth of Sunrise River at St. Croix River, Chisago Co., Minn., 6-VII-76.
D. A. Etnier (4 larvae).

Distribution

     Published distribution records are available for Illinois, Michigan, and
Tennessee.  The above records extend its range into Alabama and Minnesota.

Biology

     Ross (1944) encountered this species in abundance only along the Middle
Fork and Salt Fork rivers near Oakwood, 111., where it emerged from April
through August.  Sunrise River is about 30 m wide at its mouth, and is a
clear, shallow stream with gravel substrates.  It appears to have a very
diverse fauna.  The Little Cahaba River, Bibb Co., Ala., is a pristine stream
about 25 m wide with gravel and bedrock substrates in riffle areas.  It
contains a diverse assemblage of fishes including a health population of the
threatened goldline darter (Pevo-ina awPO~Linea.ta Suttkus and Ramsey).  Cahaba
River at the Bibb Co. Hwy 27 bridge has apparently recovered somewhat from
industrial, domestic, and strip mining pollution, and also has a diverse fish
fauna including the best remaining population of the undescribed Cahaba
shiner, an endangered species.  This suggests that H. fvisoni is a small,
warm water rivers that may be intolerant of habitat alteration.

                     Hydropsyehe miss-Lss'i.ppiensis Flint
                                  (Fig. 45)

     Sydropsyohe m-tssissipp'lensis Flint, 1972.  J. Ga. Ent. Soc., 7:80 (Type
locality:  Ms., Wayne Co., Waynesboro, 2-VIII-69.  C. Bryson).

Description

     Head capsule length, 1.50 to 1.60 mm; head capsule width, 1.40 to 1.50
mm.  Frontoclypeus brown with two, large, central, yellow spots.  Numerous
bristle-like setae on posterior half of frontoclypeus.  Posterodorsal sec-
tions of genae brown with numerous oval-shaped, small, yellow spots.  Area
around eye and ventral surface of genae yellow.  Genae with numerous, bristle-
like setae.  Mandibles yellow, edged in dark brown.  Labrum yellow with many
black setae; laterally fringed with many, long yellow setae.  Nota light
brown; each with black, bristle-like setae; less numerous on metanotum.
Prosternum yellow, posterior margin black.  Prosternal and mesosternal plates
yellow, not heavily sclerotized.  All legs yellow with numerous, black setae
and small, brown spine-like setae.  Abdomen beige; scale hairs on dorsum
least abundant on segment I, increasing posteriorly.  Minute spines on dorsum
of all abdominal segments and anal legs; anal legs lacking large, heavily
sclerotized spine-like setae; covered with numerous,, short, black hairs.

Diagnosis

     The larvae of H. miss-Lssippi-ens'is most nearly resembles those of H.
sitoulans and H. •ineormoda to which it is closely related.  The color pattern

                                     95

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of the head is nearly identical in all three species, although E. mississ-ip-
p-iens-Cs can be readily separated from the other two on the basis of bristle-
like setae (Fig. 13) present on the posterior half of the f rontoclypeus .
Eydropsysks s-imulans and E. inaarmoda lack such setae on the posterior
section of the f rontoclypeus.

Material Examined      *

Mississippi:
     Leaf R., on 1-59, just S. of Laurel, Jones Co., Ms., 25-V-76.  G. A.
     Schuster, D. A. Etnier (1 male mmt,, 2 larvae).

Distribution

     This species was previously known only from the type material Flint
C1972) .

Biology

     The Leaf River, from which the pupa and larvae were collected, is a
medium-sized river with sand and bedrock bottom.  The specimens were col-
lected in the shallow riffle areas where sections of the bedrock were
partially exposed.

     Little is known of the emergence patterns of S. mLss-issi-ppiens'i.s , but
it appears that the emergence period ranges from May to August.

                                               Banks
                         Eydyopsyohe
                                  (Fig. 46)
     Hydpopsyche venulc&is Banks, 1914.
Washington, D, C., June 22).
                                         Can. Ent., 46:252  (Type locality:
     Eydpopsyahe venulcafisi  Betten, 1926.  Mem. Cornell Ag. Exp. Stat.,
101:524; Betten, 1934.  Caddis Flies N.Y. St., p. 191-192; Milne, 1936.
Stud. N. Am. Trich., 3:73 (as syn. of soalaris}; Ross, 1938.  Psyche, 45:19
(description of lectotype); Ross, 1944.  111. Nat, Hist. Surv. Bui., 23:294;
Etnier, 1973.  J. Ga. Ent. Soc., 8:273; Resh, 1975.  Trans. Ky. Acad. Sci.,
36:12.

Description

     Head capsule length, 1.65 to 1.85 mm; head capsule width, 1.45 to 1.65
mm.  Head bronze in color*  Frontoclypeus with two pairs of yellow spots
anterolaterally; anterior spots larger than posterior spots.  Spots often
fused to form two, large, anterolateral spots.  Two, large, posterior, sub-
rectangular spots on top of genae.  Laterally behind eye, bronze with many
small, yellow spots on four or five rows.  Ventrally, head bronze with some
yellow areas, especially along gular suture.  Top and side of genae with
anterolateral corners.  Posterior half of frontoclypeus with minute spine-
like setae located in yellow pockets.  Labrum brown with numerous, brown
setae, fringed laterally with numerous, yellow setae.  Mandibles golden and
                                     96

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edged in dark brown *  Nota brown; pronotum and mesonotum with numerous,
bristle-like setae; metanotum lacking such setae.  Each notum bordered
laterally with black.  Legs similarly colored as nota and with numerous,
small, spine-like setae and longer hair-like setae, not unlike other species.
Prosternum yellow, bordered anteriorly and posteriorly in black; prosternal
plates and mesosternal plates brown.  Dorsum of abdomen with numerous, scale-
l£ke setae on all segments; minute spines on dorsum of abdominal segments I -
VII.  Minute spines present on venter of anal legs and venter of last two
abdominal segments; large, heavily sclerotized spine-like setae on venter of
anal legs lacking.

Variation

     The major variation found in this species involves the spots on the
frontoclypeus.  There are typically two pairs of these spots; however, they
often fuse to form two, large, diagonal, yellow spots anterolaterally on the
selerite.

Diagnosis

     Hycfropsyehe venula.'pis is readily identified on the basis of head capsule
coloration.  There are typically two, large subrectangular, yellow areas on
the top and posterior portion of the head (Fig. 46).  The frontoclypeus has
two pairs of anterolateral spots which may fuse into two, large, lateral
spots.  Additional characters which may aid in identification are the presence
©f minute spines on the dorsum of the abdominal segments I - VII and small
spine-like setae on the posterior half of the frontoclypeus (Fig. 15A and B)
situated in yellow, round pockets.

Material Examined

North Carolina:
     Tuckaseegee R., at U.S. 411, Dillsboro, Jackson Co., N.C., 27-VI-76.  D.
     A. Etnier (2 male mmts.).

Tennessee:
     Little R., 1.0 mi. S. of Townsend, Blount Co., Tn., 27-VIII-74.  M.
     Hughes, B. Smith, G. A. Schuster (2 male mmts., 4 larvae); Conasauga R,,
     at Tn. 74 bridge, Bradley Co., Tn.; 4-V-75.   C, and G. A.  Schuster,
     Etnier Family (3 male mmts., many larvae); Little R., 4.0  mi.  S, of jet.
     River Rd. and U.S. 411, Blount Co., Tn«s 23-IV-75.  C. and G.  A. Schuster
     (1 male mmts.); Little R., 1.5 mi. S. of jet. of River Rd, and U.S. 411,
     Blount Co., Tn., 23-IV-75.  C. and G. A. Schuster (10 male and 4 female
     mmts., many larvae); Little R,, at U.S. 411, Blount Co., Tn.,  23-IV-75.
     C. A. and G. A.  Schuster (11 male mmts., several larvae);  Little Tennes-
     see R., at Coytee Springs;, Loudon Co.,  Tn.,  6-V-75.  W. C. Starnes, G.
     A. Schuster (several male and female mmts.); Little R,, 2.0 mi. S.  of
     Townsend, Blount Co., Tn., on Tn. 73, 14-VII-76.   M. Hughes, G. Schuster,
     W. Dickinson (several larvae); Little R.,  at Alcoa Water Treatment Plant,
     0.5 mi. E. of Tn. 33, Blount Co., Tn.,  22-VII-75.  G. A. Schuster (1 male
     mmt., several larvae); Nolichucky R., at Solomon Island, 12.6  mi. N. of
     Newport, Greene and Cocke Co. line, 26-X-75.  D,  A. Etnier (4  larvae).

                                     97

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Virginia:
     Little R., at Girls Camp, Giles Co., Va., 29-V-75.  D. A. Etnier, G. A.
     Schuster (2 male mmts., 5 larvae); Stoney Cr., at jet. of Co. Rd. 80 and
     Va. 122, Bedford Co., Va., 30-V1II-76.  D. A. Etnier, G. A. Schuster
     Cseveral mmts. and larvae).

Distribution

     Published distribution records include the following states:  Kentucky,
Missouri, New York, Tennessee, Virginia, Washington D. C., and Wisconsin.
New state records given above are from North Carolina.

Biology

     Hydropsyche venulopis occurs in medium-size rivers characterized by
large riffle areas.  They are typically found on medium-sized rocks covered
abundantly with aquatic vegetation.  The pupae are often attached to the
vegetation, rather than the rock.  In this situation the pupal case is
completely cylindrical and enclosed on all sides by small pebbles.  Hydro-
psyche venul,ca"i>s is often collected in association with Sympkitopsyohe morosa,
8. sparna, and S. l>vonba.

     This species has an emergence period from early April to late September.

                          Hydropsyche hoffmani Ross
                                (Fig. 16, 47)

     Hydropsy die hoffmani Ross, 1962.  Ent. News, 73:129-130  (Type locality:
Radford Arsenal, Montgomery Co., Va., 4-10-VIII-56.  R. L. Hoffman).

Description

     Head capsule length, 1.85 to 2.0 mm; width, 1.70  to 1.80 mm.  Head
ventrolaterally yellow; some dark pigmentation may be  present on stridulabory
surfaces.  Except for area around the gular suture posterodorsal part of head
yellow;  area around suture dark brown.  Area of genae  bordering suture of
genae and frontoclypeus .dark brown.  Genae also with wide, transverse, brown
band behind eye.  Posterior half of frontoclypeus uniformly dark brown.
Frontoclypeus with a pair of small but distinct yellow spots  in the middle
of the sclerite.  Just posterior to apical margin of frontoclypeus and
anterior to the paired spots is large, transverse, subrectangular, yellow
area.  Apical margin broadly convexly rounded and trimmed  in  dark brown.
Frontoclypeus with a distinctly raised semicircular ridge  originating near
the anterolateral corners and running just posterior to tentorial pits.
Entire head shiny with very few hairs or  spines on the dorsolateral aspects
of the genae.  Those present are very small,  clear, and inconspicuous.  Head
with a naked appearance, except under higher magnification.   Body of  labrum
golden covered with long, black setae; anterior margin black; laterally
fringed  with tuft of long, yellow hairs.  Mandibles dark brown.  Anterior
ventral  apotome with large, rounded, and  distinct tubercle.   Thoracic nota
golden with very few spine-like setae and covered with thin,  short, brown
hairs.   All legs brown with first three  segments  trimmed  in black.  Segments

                                     98

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  of legs abundantly covered with distinct,  brown spine-like setae.   Prosternum
  mostly yellow.   Poststernal plates solidly sclerotized.   Abdomen brown to
:. yellowish green with numerous,  short,  black hairs.   Dorsum of abdomen covered
  with scale hairs with greatest  numbers on  posterior segments,  fewest  on
  segments I - III.  Interspersed between the black hairs  and scale  hairs on
  segments I - III of the abdomen are minute brown spines;  none posterior to
  segment III.  Dorsal sclerite of anal  legs covered  with  black setae;  ventral
  aspects of anal legs covered by numerous,  heavy,  dark brown spine-like setae.

  Variation                                       .-..,..


       The only variation discovered in  the  numerous  larvae examined is  that of
  head coloration.   Some larvae are distinctly darker in pigmentation so that
  the large,  yellow area at  the anterior section  of the  frontoclypeus is quite
  small and inundated with dark brown pigment.  The same may be  true with the
  large,  yellow areas on the lateral aspect  of the  genae.

  Diagnosis

       This unique  and extremely  interesting  species may be  separated from other
  sca^ca>^s  group  species  in  a number  of ways.  First,  the color pattern  of the
  head  and  nota is  unique in Eastern  North American HydvopsyoTie.  Second,  the
  almost  complete lack of setae on  the head and nota which give the larva a
  clean  shaven" appearance  is unique to  this species.   Third, the frontoclypeus
  is  convexly rounded  anteriorly  and  has very distinct semicircular ridge
  running from the  anterolateral  corners  to just posterior to the tentorial pits.
  Fourth, the mesal portion of the anterior ventral apotome is produced  into a
  prominent and rounded tubercle.

 Material Examined

 Virginia:
      Roanoke R., off U.S. 11 and 460,  just outside of ;Salem, Roanoke Co., Va.,
      28-V-75.  G. A. Schuster, D. A. Etnier (several larvae); Roanoke R., at
      wayside park on U.S. 11, about 10 mi.  W. of Salem, Roanoke Co., Va.,  •
      5-VII-75.  D. A. Etnier (several  larvae);  Roanoke R., at Va. 419 in
      Salem, Roanoke Co., Va., 10-VII-76.  G. A.  Schuster (very many larvae,
      many male and female rnmts.).   •                   ,

 Distribution

      The above Roanoke River localities are less than 60  miles .from the New
 River at Radford Arsenal, the,.type locality and  only previous record of this
 species.

 Biology                     •        , ,          .

      The Roanoke River, from which all  larval and metamorphotype material of
 S.  koffmani was  collected,  is a  medium-sized river.   At the Va.  Hwy. 419 lo-
 cality,  the river is about  6.0 to 10 meters wide and 0.3  to 1.0 meters  deep.
 The current is slow to moderately fast,  and the  water quality appears  good.
 The shore vegetation consists of shrubs and mixed  deciduous trees which give

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good shelter.  The bottom is variable, consisting of small rocks and gravel
in the riffle areas and large rocks in the long, smooth runs of the river
where the water is the deepest.

     Two species of Hydropsyohe were collected at this locality, and two from
Symphitopsyche.  The two Symphteopsyahe species, S. walk&ri and S. bronta,
were restricted to shallow riffle areas of the river where they were^found on
medium-to-small-size rocks.  The two Hydropsyohe species, H. hoffmani and
E. leonardi, were collected only in the deeper and faster runs of the river.
None were taken in the shallow riffles with S. bvonta and S. walkeri.

     There seemed to be a definite preference for very large rocks  (approxi-
mately 45 kilograms) with large, flat areas, a situation quite unusual for
Hydropsyche larvae.  Most1 species prefer rocks with many crevices and cracks
in which to anchor and build larval retreats.  It was found that H. hoffmam
preferred the upper areas of these rocks and that most of the pupal cases
were attached to the flat surfaces on top of the rocks.  In contrast, H.
leonardi was collected closer to the substrate.  Hydropsyohe hoffmani, and
H. leonardi larvae were never found in larval retreats close to one another.
It was found that the most effective collecting technique for these  two
species was to slowly feel the surfaces of the rocks underwater for  retreats
and pupal cases since the strength of the current  and the weight  of  the rocks
prohibited removal of the rocks for examination.

     Limited emergence data are available for this species.  From the collec-
tion dates of the type material and the metamorphotypes  collected,  emergence
appears to take place at least from late June to August.  When  this  species
is more thoroughly studied, it will be found that  emergence takes place,
undoubtedly, over a much longer period of time.

                          Rydropsyehe leonccedi  Ross
                         :      (Fig. 48A and B)

     EydropsyoUe  leonardt Ross, 1938a.  111.  Nat.  Hist.  Surv. Bui.,  21:145-
 146  (Type locality:  Lovells,  Crawford Co., Mi.,  2-V-36,  along  N. branch
AuSable R.,  2 mi. above town.   J. W.  Leonard).

     Hydropsydhe  leanxedi:   Ross,  1944.   111. Nat. Hist.  Bui.,  23:294;
Leonard  and Leonard,  1949.   Occ.  Pap. Mus.  Zool.,  Mich., 522:9.

 Description

     Head capsule length,  1.80 to 1.95 mm;  head capsule width,  1.50 to  1.65
 mm.  Head on top  and bottom dark brown to black;, lateral aspects of head ^wrth
 duck-shaped mark;  large,  yellow area around eye;-:narrow, yellow baad behind
 eye expanding dorsally at back of head.   Frontoclypeus with pair of pinhead-
 size,  yellow spots posterior to tentorial pits: spots often indistinct.
 Dorsolateral aspects of genae with many,  stout, bristle-like,  black setae;
 frontoclypeus with many such setae over entire sclerite except for shiny
 central area around tentorial pits; setae on frontoclypeus as long as tnose
 on genae.  Labrum black with numerous,  black setae; fringed laterally with
 numerous, long, yellow setae.  Mandibles dark brown to black.  Nota dark

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ibrown;  pronotum slightly darker than other  nota.  All  three  nota with short,
black,  bristle-like  setae, most numerous  on pronotum,  decreasing on mesonotum,
fewest  on metanotum.   Prosternum brown, edged  anteriorly and posteriorly  in
"black.  Prosternal and mesosternal  plates brown and  fairly well sclerotized.
Legs brown,  anterior pair darker brown, almost black.  Each  leg adorned with
numerous, longer black setae and shorter  spine-like  setae.   Abdomen brown,
with numerous  scale hairs on dorsum of posterior abdominal segments;  decreas-
ing in  number  anteriorly.  Minute spines  on the dorsum of all abdominal
segments; decreasing in  number  and  size posteriorly.   Minute spines on venter
of last two  abdominal  segments  and  anal legs very small  and  inconspicuous.
Venter  of anal legs with many,  black setae.

Variation

     The variation observed in  this  species  is restricted primarily to the
pair of spots  on the f rontoclypeus .   In some specimens these spots  are dis-
tinct and easily viewed, while  in other specimens the  spots  may be  almost
totally obscured.

Diagnosis

     This species is most closely related to H« hageni and E. hoffmani.  It
may be  confused with H.  hageni  since  the color  patterns  of the head of these
two species  are  similar.  Eydcopsyche hageni (Fig- 49A and B) and H* .leonavdi
(Fig. 48A and  B) may be  separated on  the basis  of the  following characters.
First, H. leonardi possesses many black, bristle-like  setae  (Fig. 15A and B)
everywhere on  the f rontoclypeus,  excluding 'a central area around the tentorial
pits.  The setae on the  f rontoclypeus are as long and  as  conspicuous as those
on the genae, while the  f rontoclypeus of H.  hageni is  not covered with as many
bristle-like setae as H.  leonardi.  The setae present  in H.   hageni are much
smaller and rather inconspicuous; because of this lack of setal covering, the
entire sclerite has a  shiny appearance.  The genae also do not have as many
setae as those found on £T. leonca>di.  Second, the minute spines on the venter
of the last two abdominal segments and especially the  anal legs are much
larger, more numerous, and conspicuous in E. hageni than in E. leonardi.
Third, the head of 'E.  leonardi from dorsal view is subrectangular and slightly
narrower anteriorly.  The head of E. hageni, oh the other hand, is almost
square in outline from dorsal view.   Last, the head pattern of H.  leonavdi is
much darker and the pair of spots centrally located on the frontoclypeus  are
often very small and indistinct.  The ventral surfaces of the head are almost
uniformly dark brown; whereas,  the venter of H. hageni has large,  yellow areas.

Material
Virginia :
     Roanoke R., at Va. 419, Salem, Roanoke Co., Va., 5-VII-75.  D. A. Etnier
     QL male mmt., 1 larva); Roanoke R. , at Va. 419, Salem, Roanoke Co., Va. ,
     lCMVi:X-75.  G. A. Schuster (7 mmts., 8 larvae).
                                     101

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Distribution

     The only published records for this species are from Michigan; therefore,
the above collections represent a substantial increase in the known range of.
#. leonaxdt.  The U.S.N.M. also has collections of H. leonardt from the     ;
Shenandoah River near Woodstock, Virginia, which were collected by Dr. 0. S.p>
Flint, Jr.                    ......                                      -•:'
                                                                            ,-j.
Biology                                                                      ;

     Hydropsydhe leonardi is a little known species that has rarely been col--i
lected since its description.  The larvae and the metamorphotypes have been ;;
collected only from the Roanoke River in Salem, Virginia.  The Roanoke River,,r.:
at this locality is approximately 6.0 to 10.0 meters wide and 0.3 to 1.0 meter
deep.  The current is moderate to fast and the bottom consists of small rocks:
and gravel in the riffle areas and large rocks in the long, smooth runs.     ;

     Symphitopsyche walker, S. bronta, H. hoffinam-, and a depz>avata group
species were collected in the same area as H. leonavdi.  The depravata group
species, S. waVkexi. and S. lovonba were collected in the shallow riffle areas
on and under small rocks.  Hydropsyche tioffmani and H. leonopd-L were collected
in the deeper, long runs on very large rocks.  Neither H. hoffmani nor H.
leonOPd'fc were collected in the shallow riffle areas.  On the large rocks,
H. "hoffmani- was far more abundant than H. leonardi and was found mostly on
top of the rocks, while H. leonardi — both larvae and pupae — were collected
on the sides of these rocks.

     The few available records indicate that this species emerges at least
from May through July.

                          Rydropsyohe bagenl Banks
                              (Fig. 49A and B)                           :

     Hydvopsyche hageni Banks, 1905.  Trans. Amer. Ent. Soc., 32:14 (Type
locality:  "Falls Church, Va.").

     Hydropscyhe hagenl:  Banks, 1907, Cat. Neur. Ins. U.S., p. 47; Ulmer,
1907.  Gen. Ins., 60:171; Banks, 1908.  Proc. Ent. Soc. Wash., 9:155; Banks,
1936.  Psyche, 43:127-128, 129; Milne, 1936.  Stud. N. Am. Trich., 3:73  (as
syn. of H. scalaris); Ross, 1938c.  Psyche, 45:17; Brimley, 1938.  Ins. N.
Carolina, p. 251; Denning, 1943.  Ent. Am. 23:109, 111, 119-121; Ross, 1944.
111. Nat. Hist. Surv. Bui., 23:87, 91, 93, 103, 294; Etnier, 1965.  Ent. .News,
76:146;: Longridge and Hilsenhoff, 1973.  Wise. Acad. Sci., Arts and Letters,
61:176; Resh, 1975.  Trans. Ky. Acad. Sci., 36:12.                      .i;

Description                                                            .'.• •

     Head capsule length, 1.70 to 1.85 mm; head capsule width, 1.55 to 1.70
mm.  Lateral aspect of head with wide, yellow band connecting yellow area
around eye and yellow area at back of head.  Stridulatory surfaces of genae
dark brown with numerous, eliptical, yellow muscle scars; mesally venter of
head with large, yellow areas.  Dorsum of head rich brown with pair of small,

                                     102

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yellow spots just mesal to tentorial pits; sometimes with additional larger
pair of anterolateral spots.  Dorsolateral aspects of head behind frontocly-
peus with numerous, eliptical, yellow muscle scars.  The very posterior part
of dorsum of head lighter than rest of head, often approaching yellow.
Dorsolateral aspects of genae with moderate number of stout, bristle-like
setae and numerous very thin and short, yellow hairs.  Frontoclypeus with
bristle-like setae only at extreme anterolateral corners.  Posterior half of
frontoclypeus with some very short, clear spine-like setae directed anteriad.
Due to lack of abundant setae covering, all of head rather shiny, especially
frontoclypeus.  Labrum dark brown with brown to black setae; laterally with
fringe of yellow setae; mandibles dark brown.  Nota brown; pronotum slightly
darker.  Each noturn with moderate number of bristle-like, black setae; pro-
notum with many, small, round, yellow spots from which emerge thin, yellow
setae like those of genae.  Numerous, short, black setae cover mesonotum and
metanotum; last two nota without numerous, small, yellow spots.  Legs light
brown to yellow with numerous spine-like setae and long, black setae.
Abdomen brown; scale hairs abundant on posterior segments, decreasing ante-
riorly, few on segment I.  Minute spines present on dorsum of all abdominal
segments decreasing in size and number posteriorly.  Numerous, black hairs
on venter of anal legs; heavy sclerotized spine-like setae lacking.  Minute
spines present on venter of last two abdominal segments and anal legs; spines
conspicuous at 100 X magnification.

Variation

     The degree of variation in E.  hageni, seems to be restricted to the color
pattern of the frontoclypeus.  Typically, there are two, small, yellow spots
mesal to the tentorial pits, but the degree of distinctness is variable;
often they are indistinct.  On occasion, a second larger pair of yellow spots
may be present anterolateral to these spots.

Diagnosis

     Hydropsyohe hageni, (Fig. 49A and B) is most closely related to H.  leonardi
(Fig. 48A and B), and the two larval forms exhibit similar coloration patterns.
However, they can be separated by a number of characters.  First, the lack of
bristle-like setae on the surface of the frontoclypeus of H.  hageni,, except in
the anterolateral corners, is evident; the frontoclypeus of H. leonardi, is
abundantly adorned with such setae.  The genae of E.  hageni. also do not have
as many bristle-like setae as are found on H.  leonardi,.  For this reason, the
head of E. hageni, especially the frontoclypeus, takes on a glossy appearance.
Second, the minute spines on the venter of the last two abdominal segments and
anal legs in S. hageni are very conspicuous and large under 100 X magnification.
They may even be observed under the high magnification of a dissecting micro-
scope.  In S. leonardi,, on the other hand, the spines are less abundant and
conspicuous at even 100 X magnification.  Third, the color of the sclerites
of the head and thorax of H.  leonardi, approach black, while the same sclerites
in H. hageni. are a rich brown.  Last, the venter of the head of E.  hageni, has
large', central, yellow areas while the venter of the head of E. leonardi, is
more or less unicolored dark brown.
                                     103

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Material Examined

Kentucky:
     Cumberland R., on Cumberland Falls Rd., 5.0 mi. W. of Ky. 264, Whitley
     Co., Ky., 25-VI-75.  G. ₯. Wolfe, G. A. Schuster (1 male mint., numerous
     larvae).

Tennessee:
     Clinch R., at Frost Ford, approx. 3.0 mi. E. of Sneedville, Hancock Co.,
     Tn., 9-IX-75.  G. A. Schuster (several mmts., many larvae).

Distribution

     Published state and provincial records include the following:  Illinois,
Kentucky, Manitoba, Maryland, Minnesota, North Carolina, Virginia, and
Wisconsin.  In addition to these is the Clinch River locality in Tennessee.

Biology

     Ross (1944) reports collecting larvae and pupae in the rapids of the
Kankakee and Rock Rivers of Illinois, both of which are fairly large rivers.
This is consistent to the type of habitat in which we have collected larvae
and pupae in the southeast.  Both the Cumberland and Clinch Rivers where E.
?iagen£ was collected are large rivers with extensive rapids areas and some
bedrock.  The larvae and pupae were taken in the fastest water on both large
7?0qks and bedrock.

     Emergence of #, 'hagen'i seems to take place from May through September,.

                   By dropsyche patera Schuster and Etnier
                                  (Fig. 50)

     Rydvopsyche patera Schuster and Etnier, 1978.  Kans. Ent. Soc. 51:218-221
(Type locality: Harpeth R., Tn.).

Description

     Head capsule length, 1.60 to 1.75 mm; head capsule width, 1.35 to 1.45
mm.  Head yellow with dark color pattern on top of head; some dark pigmenta-
tion on stridulatory surfaces on bottom of head.  Frontoclypeus with striking
color pattern; anterior half with seven, large, yellow spots; three spots in
transverse row just behind anterior margin; row of four spots behind and
parallel to  these; in second row, lateral spots may join mesal spots.
Posterior half of frontoclypeus mottled yellow and brown.  Genae with broken
brown transverse band behind eyes.  Anterior margin of frontoclypeus slightly
convex.  Genae with scattered, stout, bristle-like setae; many smaller yellow
to brown spine-like setae on posterior half of frontoclypeus and dorsolateral
aspects of genae.  Labrum tan; numerous, slender, black setae on top of
selerite; brush of long, golden setae form lateral fringe.  Mandibles yellow;
margins and  apical end dark brown.  Nota straw colored; pronotum and mesonotum
w$fch scattered bristle-like setae; bristle-like setae absent on metanotum.
     yellow; anterior legs slightly darker; segments of all legs amply adorned.

                                     104

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with long, black setae and short, yellow and brown spine-like setae.
Prosternum yellow, posterior margin black; prosternal and tnesosternal plates
brown.  Scale hairs sparse on abdominal segments I - IV; increasing in den-
sity from segments V - VIII.  Minute spines present on dorsum of segments
I - VIII; present on venter of segments VIII - IX and on anal legs.  Venter
of anal legs with large, golden spine-like setae similar to those on ventral
patches of segments VIII - IX.

Diagnosis

     This species may be recognized on the basis of the following combination
of characters.  First, the color pattern of the head is unique in Eastern
North American Hydropsyahe.  This species exhibits the most striking color
pattern of any known species in the saalca"i,s group (Fig. 50).  Second, large,
golden spine-like setae are present on the venter of the anal legs.  Third,
the scale hairs are sprase (Fig. 11) on segments I - IV of the abdomen.

Material Examined

Tennessee:
     Harpeth R., 1.3 mi. N of jet. U.S. 70 and Co. Rd. 7338, Cheatham Co.,
     Tn., 2-V-75.  D. A. Etnier, G. A. Schuster (12 male, 2 female mmts.,
     very many larvae); Harpeth R., 1.3 mi. N of jet. U.S. 70 and Co. Rd.
     7338, Cheatham Co., Tn., 10-IX-75.  G. A. Schuster, D. A. Etnier, R.
     D. Suttkus, M. H. Hughes, G. W. Wolfe, W. C. Starnes (several male
     and female mmts., many larvae).

Distribution

     This species is known only from the Harpeth River in Cheatham County,
Tennessee.

Biology

     This species has been collected only on two occasions, and, therefore,
very little is known concerning its biology.  The habitat in which it was
collected is a medium warm-water river which serves as a basin for a great
deal of agricultural runoff.  The river in these stretches contains a
moderate-to-heavy silt and suspended organic material load.  The bottom
consists of sand, coarse gravel, and large boulder areas.  The boulders lie
in the swiftest rapids and serve as the habitat for this species.  The larvae
and pupae were mostly collected by running the fingers across the surfaces of
these rocks and feeling for the larval retreats and pupal cases.  Many, .of the
boulders have a lush growth of aquatic vegetation occurring as long strands.
Most of the pupal cases were attached and well concealed in this vegetation.
Only one other Eydropsyohe species was collected at this locality, and it was
found to be living in close proximity to E. patera. ,  This species is H.
s-imulans, which was not as abundant as^H. patera.

     Since mature pupae were collected,in both May and September with instars
in various stages of development, it ;is surmised that the emergence period
for this species is from late April t:o .late September.
                                     105

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Remarks

     Edwards (1966) reported H. plaooda from the Stones River in Davidson
County, Tennessee.  In correspondence with him, he made the following remarks
concerning these specimens:  "I did compare most of my material with types in
Ross1 collections at Urbana, Illinois, and both he and I did note that the
forms I was referring to as H. plaooda were not precise fits with his."
Unfortunately, these specimens have been lost, and the determination cannot
be substantiated.  However, we feel certain that those specimens Edwards
called H. plaooda were in actuality Eydropsyahe patera, since his collections
were only a few miles from the above Harpeth River locality.  To complicate
matters further, Edwards' Stones Rivers locality is now impounded.

     Hydropsyche plaooda has not yet' been associated, and because HydropsyoUe
patera is closely allied to it, it is suspected that its larvae is not too
unlike that of Hydropsyolie patera.  Judging from published records, H.
plaooda is a very-large-river species.  This may be part of the reason for
it remaining unassociated since larger rivers are difficult to collect.
                                     106

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 	•   1924.  Morphology of the prolegs of Trichopterous larvae.  Ann.  Entv
     Soc. Am., 17:97-102.

 Leonard, Justin W., and Fannie A.  Leonard.  1949.   An  annotated  list of   />;
     Michigan Trichoptera.  Occ. Pap.  Mus.  Zool.  Mich.,  522:35 pp.

 Lepneva, S.  G.  1964.   Larvae and pupae of Annulipalpia,  Trichoptera.   Fauna
     of  the U.S.S.R.  Zool.  Inst.  Akad.  Nauk.  S.S.S.R.,  new Ser.,  88:1-638
     (Trans.  Isreal  Program Sci. Trans.,  Inc.,  1970).

         1966.  Larvae and pupae of Integripalpia,   Trichoptera.   Fauna of  the
     U.S.S.R.   Zool.  Inst.  Akad.  Nauk.  S.S.S.R.,  new Ser.,  95:1-560.   (Trans.'
     Isreal Program Sci.  Trans.,  Inc.,  1971).

 Lloyd,  J.  T.   1915.   Wood-boring trichoptera.  Psyche,  22:17-21.

         1921.   The biology of North American caddis fly larvae.  Lloyd Library
     of Botany,  Pharmacy and Materia Medica Bui.,  21:1-124.

 Longridge,  J. H.,  and W.  L.  Hilsenhoff.   1973.  Annotated list  of Trichoptera
     (caddis flies)  in Wisconsin.  Wise. Acad. Sci., Arts and Letters,  61:
     173-183.

 Marshall, A. C.  1939.   A qualitative and quantitative  study of the Trichoptera
     of western  Lake Erie (as indicated by light trap material).  Ann.  Ent. Soc.
     Am., 32:665-688.

 McLachlan,  R.   1863.   Notes  on North American Phryganiedae, with special
     reference to those contained  in the collection of the British Museum.
     Entomologist's  Annual,  1863:155-163.

 Merrill, D., and G.  B.  Wiggins.   1971.  The larva and pupa of the caddisfly
     genus Setodes  in North America.   Life Sci. Occ. Pop., R. Ont. Mus.,
     19:1-12.                                                            i

 Milne,  D. J.  1943.   The distribution and life histories of the caddis flies
     of Waskesiu Lake,  Saskatchewan.   Can.  Ent., 75:191-198.

 Milne,  Lorus J.  1934-1936.  Studies  in North American Trichoptera, I, II,
     III.  Cambridge.                  ,                                ,

Milne, Margery J.  1938.  The "Metamorphotype Method" in Trichoptera.   J.  N.Y.
     Ent. Soc., 46:435-437.                                             -".

	•  1939.  Immature North American Trichoptera.  Psyche, 46:9-19.

Morse, W. J., and R. L. Blickle.  1953.  A check list of the Trichoptera
     (caddis flies)  of New Hampshire.  Ent. News, 64:68-73,  97-102.
                                     112

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Muttkowski,  R. A.   1929.   The  ecology of  trout  streams  in Yellowstone National
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Neave, F.  1934.  A contribution to the aquatic  insect  fauna of Lake Winnipeg.
     Int. Rev. Hydrobiol.,  31:157-170.

Nimmo, Andrew P.  1966.  A list of the Trichoptera taken at Montreal and
     Chambly, Quebec, with  descriptions of three  new species.  Can. Ent.,
     98:688.

	.  1971.  The  adult Rhyacophilidae and Limnephilidae (Trichoptera) of
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nt:  '
Orcutt, A. W.  1934.  The larvae.  In C. Betten, The caddis flies or Trichop-
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Packard, A. S.  1876.  Guide to study of insects.  New York.

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Resh, V. H.  1973.  Biology and immature stages of caddis flies of  the genus
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	.  1975.  A distributional study of the caddisflies of Kentucky.  Trans.
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	'_•  1938b.  Lectotypes of North American caddis flies in the Museum, of
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                                     113

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                                     114

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Wiggins, G. B.  1960.  A preliminary systematic study of the North American
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	.  1963.  Larvae and pupae of two North;American L imnephilid caddisfly
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	.  1965.  Additions and revisions to the genera of North American caddis-
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	.. 1973a.  A contribution to the biology of caddisflies (Trichoptera) in
     temporary pools.  Life Sci. Occ. Pap., R. Ont.  Mus., 88:1-28.

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     Pap., R. Ont. Mus., 91.                       .     .    -

	.  1977.  Larvae of the North American caddisfly genera (Trichoptera).
     U. of Toronto Press, Toronto, Can.,"401 pp.              .        V"'

Wiggins, G. B., and N. H. Anderson.  1968.  Contributions to the systematics
     of the caddisfly genera Pseudostenophylax and Philooasea, with special
     reference to the immature stages (Trichoptera:Limnephilidae).  Can. J.
     Zool., 46:61-75.
                                       t (;';•'.-•'•-,      '
Yamamoto, T., and G. B. Wiggins.  1964.  A comparative study of the North
     American species in the caddisfly genus Mystaoi-des (Trichoptera:Lypto-
     ceridae).  Can. J. Zool., 43:1105-1126.

                                     115

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                                  GLOSSARY
Anal Papillae — Four elongate lobes arising from the anal opening, which may
     or may not be extruded in preserved specimens.  They function both in
     respiration and ionic transport.                                       '

Appressed Hairs - Thin, hair-like, dark brown to black  setae covering the
     membranous epithelium of the abdominal and thoracic segments.
             Setae - Stout, dark brown to black setae occurring on the
     thoracic nota and sclerites of the head, especially the genae and
     f rontoclypeus .

Carina - A keel-like structure produced in Hydropsyohe by the suture of a
     depressed frontoclypeus and elevated genae.

Club Hairs - Setae on the dorsum and sometimes the lateral aspects of the
     abdominal segments of Symphitopsyohe.  These setae are elongate, slightly
     widening at the apex and only a bit wider than the appressed hairs.
Epicranial Arm - Or frontoclypeal suture.
     genae to the frontoclypeus.
That suture which connects the
Epicranial Stem - Or coronal suture.  That suture at the posterodorsomedian
     area of the head which joins the genae or parietals.

Epicranial Suture - The entire suture consisting of right and left epicranial
     arms and epicranial stem.

Foretrochantin - The sclerite at the base of the anterior leg which in
     Hydropsyche, Symphitopsyche, Chewnatopsyehe, and occasionally in
     Potamyia is forked.

Forked Stridulator - Sensu Ross, 1944.  See foretrochantin.

Frontoclypeus - In Trichoptera, a single sclerite composed of the frons and
     clypeus.  In Hydropsyahe and Symphitopsyehe, this sclerite is subtri-
     angular in outline and separates the genae dorsally.

Genae - Or parietals.  These large sclerites constitute the greatest propor-
     tion of the head capsule laterally and ventrally, and are dorsally
     separated by the frontoclypeus.
                                    116

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 Gular Suture - Or ventral ecdysial line.   The suture which connects  the
      genae ventrally.

 Hairs - Long,  thin,  flexible setae.

 Larval Retreat - Tube-like structure,  expanded at  the opening,  in which the
      larva lives.  It  consists  of  silk Woven into  fishnet-like  mesh,  often
      with a variety  of debris attached to  the outside.

 Maxillary Palp - A lobe of the  maxillary stipes.

 Metamorphotype (mtt.)  - Or pharate adult.  That life history  stage to
      the order Trichoptera,  which  allows for absolute adult-larva association.
      It is at  that time in the  life  cycle  of the caddisfly, when  the  sloughed
      larval sclerites  have been pushed to  the rear of the  pupal case, and
      pupal metamorphosis has progressed to just prior to emergence of the
      adult.  At this time the adult  genitalia has sufficiently  sclerotized  to
      allow species identification.

 Minute  Spines  - Spines located  on  the  dorsum of some or all abdominal segments
      of Hydropsyche  species,  usually decreasing in size posteriad.  Such
      spines  are also found  on the  venter of  the last  two abdominal segments
      and anal  legs.

 Muscle  Scars - Round or elliptical spots visible on  the genae and/or frontb-1-
      clypeus which are produced by attachments of muscles  in the head.  They
      may be  either lighter  of darker than  ground color of  the sclerite.

 Fharate Adult  - See  Metamorphotype*

 Pleural Gills  - Elongate, conical  gills located on the lateral areas of
      segments  III  -  VII.  All known Eastern North American species of
      Hydropsyche and Symphitopsyche possess 1, 3, 3,  3, and 2 pleural gills,
      respectively  from segments III - VII.

 Poststernal Plates - Subtriangular plates  immediately posterior to prosternum;

 Prosternum -Strap-like  sclerite present on the venter of the prothorax.

Pupal Case - The cylindrical structure usually composed of pebbles and/or
      sand grains attached to the substrate in which pupation takes place.

Scale Hairs - Wide,  subtriariguOiar and scale-like setae found on the dorsum
      and  sometimes the lateral'-aspects of some or all abdominal segments of
     Hydropsyche species.  These hairs are usually most abundant on the last
      two or three  segments/ decreasing in number and sometimes in size
      anteriad.

Setae - Hairs, bristles, heavily sclerotized spine-like projections,  and
      spurs which arise from pits in the cuticle.
                                     117

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Spines - Extensions or processes of the cuticle.

Spine-like Setae - Setae, which are usually heavily sclerotized and pointed,
     giving the appearance of spines.

Stridulatory Surface - Ventral striated area of genae on which is rubbed a
     prominence of the front femur.

SuTmentum - Mentum, sensu Ross, 1944.  The lobed sclerite just anterior to
     the Anterior Ventral Apotome.

Tentorial Pits - External depressions near the center of the frontoclypeus
     in Hydropsyche and Symphitop'syche, that internally give rise to the
     tentorial arms which act as sites for muscle attachments.

Ventral Apotome - Or gular sclerite, sensu Ross, 1944.  This sclerite in
     Hydropsyehe and Symphitopsyche is divided into two distinct sclerites;
     the Anterior Ventral Apotome and Posterior Ventral Apotome.  The
     anterior ventral apotome (Submentum, sensu Ross, 1944) is the subtri-
     angular sclerite just posterior to the submentum.  The posterior ventral
     apotome or protogula is much reduced and located just posterior to
     posterior end of gular suture.
                                      118

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                  APPENDIX A






SPECIES ASSOCIATED FROM EACH GENUS AND SPECIES



GROUP WITH A LISTING Of STATES FROM MICH THE



 METAMORPHOTtPES AND/OR LARVAE WERE EXAMINED
                      119

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                               Genus Ey'dropsyche'
                            DepVdvata Species Group
Eydropsyche betterd-  Ross' (R)
         E. cacol-Lna Banks
         H. decalda  Ross
         E. depravata Hagen
         H. potcmaoens'Ls Flint
         H. ei-issoma Ross
Ky., N.C.> Tn., Va.
N.G.
Ga., La*, Tx.
Tn., Ga.
Va.
Ga.
          (R)  « Originally associated by H. H. Rosss  1944.
                                       120

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                                   s Species Group
Hydropsyehe aevata ROSS  (R)




         H. bidens Ross



         H. demora Ross




         H. dioantha, Ross  '



         H. Tiageni Banks




         H. hoffmani Ross




         H. inaormoda Hagen




         H, leonapdi Ross



         H. m-issis&ppiens-is Flint



         H. OTT-ls Ross (R)



         H. phalerata Hagen (R)



         H. soalar-i-s Hagen



         H. simulans Ross (R)




         H. valanis  Banks



         H. venutccc-ls Bain.ks




         H. pateva Scnuster and
II.




Mn,, II.,




Tn.




Ky., Mn., Oh.




Ky., Tn,, Va.




Va.




II.




Va.




Ms.




Mn,, Oh.




Ky,, Tn., Va.




Mn., Tn., Va,,  Wi,




II,, Ky., Mn.,  Tn.,  Tx,




Oh,




N.C., Tn., Va.




Tn,
          (R)  = Originally associated by H, H. Ross,  1944.
                                      121

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Hyfoopsyche Species Only Associated by Ross  (1944)
Eydropsyohe oucavis Ross
            A.  Cuan^s Species

                                   II-
           B, Scc&aris  Species
Hydropsyche arinale Ross



         H. frisoni Ross
                                   11.



                                   II •
                        122

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                             Genus SymphZtopsyche
Symphitopsyche alhedra  (Ross)




            S. b-Lfida  (Banks)  (R)



            S, bvonta  (Ross)  (R)




            S. oheilonis  (Ross)  (R)



            5. maoleodi  (Flint)




            S. morosa  (Hagen)




            5. piatrix  (Ross)   **




            S. reffurvata  (Banks)  (R)



            S. viola  (Banks)



            S. slossonae  (Banks)  (R)



            S. spama  (Ross)



            5. Ventura  (Ross)



            S. walker-L  (Betten & Mosely)



            S, etnieri  (Schuster  & Talak)
N,C., Tn.




Mn., Wiv-




N.C., Tn., Va.




Oh., Tn., Va.




Ga., Tn.




Mn., N.C., Tn., Va.




Ar., Mo.




Mn., Wi.




Mn.




Mn., N.C., Tn., Va.




Al., Ky., N.C., S.C., Tn., Wi.




Tn.




Mn., Va.




Tn.
             (R) = Originally associated by H.  H.  Ross, 1944.




             **  = Probable Association
                                     123

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                    APPENDIX B






UNASSOCIATED VIDROPSICHE AND SyMPHITOPSXCHE SPECIES




     WITH LISTING OF ALL LITERATURE AND KNOWN




       DISTRIBUTION OF EASTERN NORTH AMERICA    .  ,
                                   io
                        124

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                         HYDROPSYCiSE ALVATA DENNING
Denning, 1949c.  Brook. Ent, Soc., Bui.,
44:40 (Type locality:  Jackson, Ms,,
24-30-IV-46.  P. H. Harden).            ..-..:-,  ,Ms,, II.

Unzicker et al,, 1970.  J. Ga. Ent. Soc.,   .
5:171.                                         Ak.
                        HYDROPSYCHE BIDENTATA DENNING
Denning, 1947d.  Ent, News, 58:249  (Type
locality:  Columbia, S.C,, 5-VI1I-43.
D. G. Denning).
S.C.
                          BXDROPSXCBE CA.TA.mA. ROSS
Ross, 1939a.  Proc. Ent. Soc. Wash.,
41:67-68  (Type locality:  Catawba R,,
Catawba, N.C., 23-IV-38.  Ross and Burks).     N.C.

Brimley, 1942.  Ins. N.C. Suppl., p. 15,       N,C.

Ross, 1944.  111. Nat. Hist. Surv. Bui,,
23:294.                                        N.C,
                                      FATTIGI ROSS
Ross, 1941a.  Am, Ent. Soc., 67:88-89
(Type locality:  6 mi. W- of Concord, Ga. ,
ll-V-39.  P. W. Fattig).                       Ga.

Ross, 1944.  111. Nat, Hist. Surv. Bui.,
23:294.                                        Ga,
                                     125

-------
Denning, 1948b,  Ann,  Ent;,.  §o
-------
\mwomGm
                                              mm
 Ross,  1941.   Trans. Am.  But. Sos,, 67;87*T
 88 (Type locality:  Namakagon R, ,
 Wi.,  5,  6-VI-36,  Frispn and Rpss,)

 Denning,  1943.  Ent. Amf , 23:109, 111,
 XX 0 *                                  '
 Ross,  1944.   111. Nat. Hist,
 23:91,  93, 103-104, 294,
 Etnier,  1965.  Ent. News, 76:146,
 Edwards, 1966.  J. Tenn, Acad,
 41:120.
Nimmo, 1966.  Can. Ent., 98:691r

Corbet et al.  Can, Ent,, 98 a288, 3,293- ,

Longridge and Hilsennpff, 3,9 73 f . . Wise.
Acad. Sci., Arts and Letters. ? 6
Ross, 1947a.  Trans. Am, Ent.
73:139  (Type locality:
College, Greene Co., Tn,, 8-VIII-46,
Wright).
Ross, 1938b.  Ill, Nat. Histi
Bui., 21:148-149 (Type locality;
Bloomer, Wi., 5-VIH36,  Prison
Ross) .
Denning, 1943.  Ent, Am,, 23:110,
112, 124-125.            
-------
Morse and Blickle, 1953.  Ent. News,
64:71.

Etnier, 1965.  Ent. News, 76:146.

Blickle and Morse, 1966.  Me. Ag. Exp,
Sta. Tech. Bui., 24:6.

Nimmo, 1966.  Can. Ent., 98:691.

Corbet et al., 1966.  Can. Ent.,
98:1291.

Longridge and Hilsenhoff, 1973.  Wise.
Acad. Sci., Arts and Letters, 61:176.
N.H.

Mn.


Me.

Quebec


Quebec


Wi.
                                    128

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                                   TECHNICAL REPORT DATA
                            (Please read Instructions on the reverse before completing)
 I. REPORT NO.
 EPA-600/4-78-060
                                                         3. RECIPIENT'S ACCESSION-NO.
 VffiMSf %BRr'1rllE IDENTIFICATION OF THE LARVAE OF THE
 CADDISFLY GENERA HYDROPSYCHE PICTET AND SYMPHlIOPSYGHE
 ULMER IN EASTERN AND CENTRAL NORTH AMERICA (TRICHOPTERA:
                                                         5. REPORT DATE
                                                          October 1978  issuing date
                                                         6. PERFORMING ORGANIZATION CODE
 Guenter A.  Schuster and David A. Etnier
                                                           8. PERFORMING ORGANIZATION REPORT NO
 9. PERFORMING ORGANIZATION NAME AND ADDRESS
 State Biological Survey of Kansas, The University of
 Kansas, Lawrence,  KS, 66044 and Department  of  Zoology,
 The University of Tennessee, Knoxville, TN   37916
                                                          10. PROGRAM ELEMENT NO.
                                                                PE 1BD612 '
                                                         11. CONTRACT/GRANT NO.
 12. SPONSORING AGENCY NAME AND ADDRESS
 Environmental Monitoring and Support Lab. - CINN,  OH
 Office of Research and Development
 U.S. Environmental Protection Agency
 Cincinnati,  OH   45268
                                                         13..TYPE OF REPORT AND PERIOD COVERED
                                                                Final
                                                         14. SPONSORING AGENCY CODE

                                                               EPA/600/06
 15. SUPPLEMENTARY NOTES
 16. ABSTRACT           •                                        ~~~~^~—~~	
      Larvae  of  the caddisfly genera Hydropsyche and Symphitopsyche are among  the most
 encountered  and abundant organisms of lotic  environments in eastern North America.   Ye
 little is known of the larval stages of these genera.   Previously, the larvae of only
 12 species of which descriptions are presented here,  were known.  Descriptions of
 larvae of an additional 27 species are here  presented for the fifst time.  Presented
 here are the descriptions of the larvae of 14 of 15 nominal eastern species of the
 genus Symphitopsyche,  and 25 of 34 of the genus Hydropsyche; 18 of 24 of the  scalaris
 group, 6 of  9 of the  depravata group, and the single species of the cuanis group, H.
 cuanis Ross.  A key is provided for known larvae,  incorporating a number of characters
 previously unused in  the taxonomy of these two genera.   The key is based on ultimate
 or penultimate  larval instars since color patterns may be more variable in earlier
 instars.  Larvae of S. piatrix have not definitely been associated with metamorphotype
 however, larvae were  collected near the type locality,  and the presumed larvae of this
 species is described,  illustrated, and keyed.   A listing of all unassociated  species,
 with known distribution and literature citations,  is given.  In addition to the descrii
 tion of the  larvae, the following are presented for each of the associated species  of
 Hydropsyche  and Symphitopsyche;  known range,  notes on the biology, diagnosis,  intra-
 specific variation,  material examined, complete literature survey and synonomies,  and
 illustration  of  the  head capsule and pronotum.
 7.
                                KEY WORDS AND DOCUMENT ANALYSIS
                  DESCRIPTORS
                                              b.IDENTIFIERS/OPEN ENDED TERMS
                                                                        C.  COSATI Field/Group
 Aquatic Biology
 Freshwater biology
 Indicator species
 Benthos
 Insects
 Larvae
 Life cycles
 F.rnl ngy
                                 Taxonomy
                                                 Trichoptera
                                                 Hydropsychidae
                                                 Hydropsyche
                                                 Symphitopsyche
                                                 Caddisflies
                                 6C
                                 6F
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